apg classification for students

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PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010 1 PHYLOGENY AND EVOLUTION OF SEED PLANTS Introduction On classifications in general, and this classification in particular A phylogenetic tree of seed plants On thinking about apomorphies Summary of the system LINKS TO ORDERS AND FAMILIES On the interpretation of the text, abbreviations, etc. Back to Main Tree MAIN GROUPINGS Asterids, commelinids, core eudicots, asterid 1, asterid 2, eudicots, gymnosperms, Magnoliophyta, monocots, N-fixing clade,rosids, seed plants. Page last updated: Fri, 18 Jun 2010 15:45:06 GMT

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Classification of seed plants by phylogeny

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Page 1: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

1

PHYLOGENY AND EVOLUTION OF SEED PLANTS

Introduction

On classifications in general and this classification in particular

A phylogenetic tree of seed plants

On thinking about apomorphies

Summary of the system

LINKS TO ORDERS AND FAMILIES

On the interpretation of the text abbreviations etc

Back to Main Tree

MAIN GROUPINGS

Asterids commelinids core eudicots asterid 1 asterid

2 eudicots gymnosperms Magnoliophyta monocots N-fixing claderosids seed plants

Page last updated Fri 18 Jun 2010 154506 GMT

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

2

INTRODUCTION

This linked series of four pages is a simplified version of the main site The focus is on

the larger patterns evident in the phylogeny evolution and diversification of seed

plants and on morphological variation in general

Systematics is a profoundly historical discipline and we forget this at our peril Only

with a phylogeny can we begin to understand diversification regularities in patterns of

evolution or simply suggest individual evolutionary changes within a clade Our

recovery of that phylogeny is the recovery of evidence of a series of unique events that

comprises the history of life Although our knowledge of the major clades of seed plants

and the relationships within and between them are still somewhat in a state of flux

much of the broad outline is clear (see the Angiosperm Phylogeny Group II 2003)

Furthermore as details of phylogeny are clarified and new findings made in anatomy

morphology etc they can be rapidly integrated the Angiosperm Phylogeny Group

system that is followed here Books are out-of-date before they appear furthermore

there is no comprehensive phylogeny-based treatment of angiosperms out-of-date or

not D Soltis et al (2005b) is the closest (also take the following link to a regularly

updated and printable Angiosperm Phylogeny Poster)

These pages are for undergraduate students with an interest in botany and some

knowledge of basic plant morphology and for those with a general interest in seed plant

evolution They are made up of a much modified Apomorphies page that has been

combined with the order pages all very much changed The focus is on providing

possible apomorphies for as many clades as possible although there is much less detail

than in the Apomorphies page (much chemistry details of ovule morphology other than

general appearance most indications of base chromosome number etc have been

removed) and I have simplified the vocabulary somewhat There is also information on

the sizes and distributions of families lists of the larger genera included features by

which families can be recognised comments about the ages (to be treated with

considerable caution) and diversification of groups and about the major herbivores

pollinators and seed dispersers associated with them notes on interesting

morphological variation summaries of what is known about infrafamilial phylogenetic

relationships etc The literature cited will allow the student find the major

morphological taxonomic and phylogenetic information about each family especially

when the Characters page is also consulted Phylogenies of most orders and of some

families (eg Poaceae) are included but not maps or images for these there are links

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

3

from each family to the more detailed treatments on the individual order pages In

larger families I tend to focus on literature that deals with monophyletic groups that

include fifty or more taxa in smaller families the coverage is more detailed

I have emphasized plant families because they are the groups - admittedly partly

arbitrary as to circumscription but now for the most part monophyletic - around which

many of us organize our understanding of plant diversity I also pay attention to

groupings of families because so much progress has been made in the last decade in

particular in sorting them out Infrafamilial groups in groups like Poaceae Malvaceae

and Ericaceae are also included and these are being added to as studies become

available

But we dont want to know just about clades we want to know what makes clades

unique the synapomorphies or shared derived characters of those clades that first

appeared in their immediate ancestors However for the most part our knowledge of

synapomorphies remains poor ss we will see finding out the composition of clades is

often easier than finding the synapomorphies for the same clades (see the discussion

below) And of course knowing about synapomorphies is just one aspect of

understanding the whys and wherefores of the evolution and diversification of seed

plants our ultimate goal

ON CLASSIFICATIONS IN GENERAL AND THIS CLASSIFICATION IN PARTICULAR

On classifications in general

Classifications in the broad sense are box-in-box group-in-group or partwhole

naming devices that we use to communicate aspects of our knowledge of things in

general From this point of view there is nothing particularly special about biological

classifications apart perhaps from their sizes For any biological classification system to

be effective it must be stable universal (ie be used by a wide range of people)

comprehensive (if too many taxa are unplaced at the level of interest the classification

is of less utility) and it must enhance communication of knowledge by helping us to

relate things (Stevens 2006a for references) Phylogenetic classifications convey aspects

of our knowledge about phylogenetic relationships of organisms Thus a family is clearly

flagged as such and is a monophyletic group that can contain several genera also

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

4

flagged as such and also monophyletic but a genus can never include families Generic

family etc names are simply words we use to denote appropriate parts of phylogenies

and minimal aspects of their relationships and the irreducible aspect of relationships

that is emphasized here is monophyly A monophyletic group is one containing only and

all descendents of a common ancestor and it is often characterizable by apomorphies

that is shared derived characters that appeared in the ancestral lineage or stem clade

of that monophyletic group

Thus I am using a flagged ie ranked hierarchy for naming taxa rather than the

unranked systems that have recently been proposed (see below) The rank terminations

used (-ales -aceae etc - the flags) merely suggest relative positions of groups in the

local hierarchy If Ericaceae and Vaccinioideae are part of the same monophyletic group

the latter must refer to a clade nested within the former even if neither can necessarily

be directly compared with Polemoniaceae and Cobaeoideae (other than all being

putatively monophyletic groups) Such a flagged hierarchy is useful as a mnemonic and

communication device (eg Stevens 2006a) It improves memorization and emphasis on

families and orders as here is a didactic device - families are monophyletic units useful

in communication major units learned by biologists and others world-wide

The distinction between grouping and ranking is extremely important as is how we

interpret the latter We can both agree that there is a genus Acer yet disagree as to

whether it should be in Aceraceae or submerged in Sapindaceae Although from one

point of view this disagreement is utterly trivial it can have profound consequences if

we misunderstand the nature of the classificatory hierarchy Taxa at the same rank are

equivalent only by designation and have nothing necessarily in common (unless they

are sister taxa) other than their monophyly Rank as used here has no meaning other

than signifying a monophyletic group that includes other monophyletic groups with

appropriately subordinate rank terminations Taxa at the same rank have often been

treated incorrectly as if they were equivalent by biologists attempting to understand

evolutionary or biogeographic problems (see Bertrand et al 2006 for detailed

discussion) even if those constructing or using classifications - including Darwin (1859) -

have been explicit about the non-equivalence of taxa at the one rank (Stevens 1997) In

fact rank terminations have relatively infrequently been used by taxonomists to reflect

absolute rank although Linnaeus (at least in theory) at the level of genus and species

may be such an example (Classifications where rank is absolute taxa at the same rank

somehow being comparable entities are class hierarchies in the strict sense - Stevens

2002 2006a) It has also been suggested that taxon rank be adjusted so that rank

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

5

somehow reflects the degree of morphological differences between taxa or that taxa at

the same rank be based on similar characters or show a similar amount of distinctness

This might be possible using phenetic methods of analysis but is very difficult if ones

classification is phylogeny-based as here it could promote instability if used in taxa

where such a criterion had not previously been used and it might also inadvertently

suggest that taxa might be equivalent (for an example see Fritsch et al 2008) There

have also been proposals that rank could reflect the age of the clade (eg Hennig 1966)

with clades that have diverged by a particular time all being given the same rank Apart

from the fact that aging times of divergence of clades is still a difficult enterprise huge

disruptions to our nomenclature would result Recent suggestions which invoke the use

of age in classifications focus on providing a standardized timeclip ie a set of letters

referring to a particular geological period that could simply be added to a conventional

taxon name (Avise amp Mitchell 2007) However even such timeclips are unlikely to come

into general use soon

It would be impossible even to think about a higher-level classification such as this

without the advances in our understanding of relationships made by the phylogenetic

analyses of molecular data carried out over the last twenty years One can then

integrate the data to be found in both classical and recent morphological studies with

these phylogenies For the dramatic changes in this area see for instance the

pessimistic attitude towards orders in Davis and Heywood (1963 107-108) The most

unsatisfactory taxon in Angiosperm classification they were indefinable their

circumscription was not fixed etc Families they thought were likely to be the largest

natural unit within the mono- or dicotyledons Along the same lines almost three

quarters of the orders (4459 monofamilial orders ignored) recognised by Cronquist

(1981) are not monophyletic ie they do not contain all and only the descendents of a

common ancestor Most of those orders that are monophyletic are very small

(Zingiberales with eight families are the largest) for families on the other hand about

two thirds (189273) are monophyletic

Turning now to phylogenetic classifications and the particular classification used

here Backlund and Bremer (1998) provide a useful discussion on the principles of

phylogenetic classification that is applicable at all levels apart from species (see also

Stevens 1998 also Albach et al 2004 Entwisle amp Weston 2005 Pfeil amp Crisp 2005 etc

for examples) Backlund and Bremers main principle is that taxa that are recognised

formally should be monophyletic However this does not indicate which particular

clades we might wish to name as families genera etc and talk about in general

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

6

conversation If a well-supported hypothesis of monophyly is a necessary prerequisite

for a group to be named it is not a sufficient prerequisite (but cf the PhyloCode -

Cantino amp de Queiroz 2006 Cantino et al 2007) Not all clades need be named indeed

it would barely be practicable (or practical) to do this To decide which clades should be

named additional criteria can be invoked Other things being equal it is helpful if 1 taxa

formally recognised are easily recognizable 2 groups that are well-established in the

literature are preserved 3 the size of groups is taken into account and 4

nomenclatural changes are minimized (Backlund amp Bremer 1998) Thus numerous small

groups have little to recommend them since individually they summarize little

information and tend to clog the memory while groups that are too big may be

amorphous Somewhat similarly Godfray and Knapp (2004 p 562) note that users

want stable informative and accessible classifications that enable easy identification

(see also Simon 2008) - although invoking users without specifying those who make up

this group is not very helpful This classification is for all interested in comparative

biology hence the emphasis on monophyly Although the clades named are sometimes

difficult to characterise there are as we shall see many ways of making such a

classification accessible to all

Problems with this emphasis on monophyly may be caused by reticulation events

such as hybridization endosymbiosis and lateral gene transfer but they are unlikely to

be common confusing factors here Genera can often be pegged to above the level at

which hybridization is at all common However in Poaceae-Pooideae-Triticeae there are

some intractable problems where extremely well established common usage and the

principle of monophyly are likely to remain at odds Many genera are certainly not

monophyletic here being allopolyploids and the genera are ultimately based on the

different genomes they contain (Dewey 1984 Loumlve 1984 Barkworth 2000 for a history

of Triticeae classification Petersen et al 2006) There is also extensive reticulation

reported within Danthonioideae (Pirie et al 2009) Evidence also increases of old

hybridization events elsewhere in flowering plants that at the very least cause

discordance between relationships suggested by different genomic compartments as in

Smedmark and Anderberg (2007 Sapotaceae) and Fehrer et al (2007) Morgan et al

(2009) and Pelser et al (2008 all Asteraceae - hybridization is likely to be quite a

problem at the generic level here) and genera like Medicagoare turning out to have

highly reticulating relationships at the species level (Maureira-Butler et al 2008) Of

course there are many problems trying to apply the concept of monophyly to species

and for many - but not all - biologists strict monophyly is a less important criterion at

this level (Funk amp Omland 2003 see much of the discussion in Houmlrandl 2006)

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

7

The major endosymbiotic events that characterize the clade of which flowering

plants are a part (and gave rise to chloroplasts and mitochondria) are very ancient and

cause no problems for the student of multicellular organisms However lateral gene

transfer has been detected in a number of situations between quite unrelated

organisms (eg Bergthorsson et al 2003 - Amborella and liverworts see also below but

cf Goremykin et al 2009) and it may be particularly common in mitochondria (Sanchez-

Puerta et al 2008) Here too there are no major problems providing one is careful

such transfers do however raise all sorts of interesting biological questions (see

Richardson amp Palmer 2007 for a summary) Although there is increasing evidence for the

importance of genome duplications - hybridization is one cause of this - at various times

during the evolution of seed plants and of palaeopolyploidy events within eg the

Lauraceae and Magnoliaceae clades (Soltis et al 2009 for a summary see eg

the Characters page for further discussion) these too do not currently seem to pose

problems for the adoption of monophyly as the sine qua non of groups to be recognised

formally in this phylogenetic classification but it can make detecting orthologous genes

difficult

The accessory principles of Backlund and Bremer (1998) should be used in

combination Thus keeping the monogenericPlatanaceae separate from its sister

taxon Proteaceae is justifiable Both are much-used names that signal well supported

well defined and easily recognisable groups that have long been recognised as distinct

have several synapomorphies and do indeed look very unlike each other Combining

the two would yield a clade with few obvious apomorphies not to mention the fact

that Nelumbonaceae should by the same logic (it is also monogeneric) also be included

in the expanded family On the other hand it is difficult to justify the continued

recognition of Callitrichaceae or Hippuridaceae monophyletic and distinctive although

they may be If they were recognised several poorly characterised clades would also

have to be carved out ofPlantaginaceae in any classification that aimed to convey a

comprehensive view of the worlds flora The continued recognition

of Valerianaceae and Dipsacaceae also tends to run into this problem (see also Pfeil amp

Crisp 2005 Orthia et al 2005 Albach 2008 etc for useful practical discussions of such

matters) But there are no absolute guidelines If Podostemaceae turn out to be sister

to Hypericaceae (for references here and elsewhere in the Introduction see the

individual families) the subsequent moderate dismemberment of Clusiaceae sl is not

be too high a price to pay for the continued recognition of Podostemaceae Hence the

somewhat provisional recognition of Hypericaceae and Calophyllaceae as well as

Clusiaceae below the families can all be recognized and the name Podostemaceae in

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

8

particular is very well established In a somewhat similar situation if Lemna and its

relatives are a clade sister to most other Araceae should they be recognised as a

separate family Gymnostachys a phenetically fairly distinctive taxon as well as the less

phenetically distinct Orontioideae would have to be recognised as a separate families

(or combined as a single family) too but Araceae in a somewhat restricted sense would

be somewhat more morphologically coherent although not greatly so and not notably

distinct However it is in the very nature of such decisions to be somewhat arbitrary

and unsatisfactory hence the emphasis on consensus classifications here and on

classifications as simply being a means to an end

A useful distinction can be drawn between crown groups and stem groups The

former are monophyletic and include the extant members of a clade and their

immediate common ancestor (see the figure below) The groups characterized in this

site are such groups Thus Proteaceae here are crown group Proteaceae apomorphies

like the single carpel four-merous perianth etc being found in this common ancestor

Stem groups on the other hand include all the members of a lineage immediately after

its split from its sister group and all branches of this lineage In the case of Proteaceae

the stem group would include everything after its split from its sister group

Platanaceae Obviously most of the organisms in the lineage that terminates in crown-

group Proteaceae are unknown only a few being known as fossils and it is not known

exactly where particular apomorphies of crown group Proteaceae evolved along this

lineage

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

9

Indeed there is no reason other than convention or convenience why any group

should not be segregated into several smaller monophyletic groups or merged to

produce a larger unit we can talk about one large thing or about several smaller things

Thinking about aspects of size findings in ethnobiology and cognitive psychology can be

used to suggest that a moderate number - probably fewer than 500 - of families is a

reasonable goal at which to aim and that groupings of taxa throughout any system

should be rather small in size (eg Berlin 1992 Stevens 1994 1997) Major systems such

as those of Linnaeus and Bentham and Hooker were constructed explicitly so as to ease

the burden on the memory (Stevens 1997 2002 see also Scharf 2007) the latter in

particular ensuring that all groups in their classification were relatively small often

containing three to eight immediately subordinate taxa - but by no means all their

groups were formally named Along the same lines Burtt (1977b) suggested that the

number of names at any rank should be at most one third those at the immediately

lower rank - and monotypic taxa might not need a formal name Consistent with such

ideas a fairly broad view of families and orders is taken here whenever the constraints

of monophyly and other criteria used when constructing classifications (see above)

permit The APG system is thus a convention consistent with ideas such as those

expressed by Backlund and Bremer (1998) for similar conventions see eg the Grass

Phylogeny Working Group (2001) and for fungi Hibbett et al (2007)

There are other views Thus Takhtajan (1997) has suggested that smaller families

are more natural This is incorrect Monophyletic groups that include fewer taxa -

Takhtajans smaller families - do not necessarily have more apomorphies than larger

groups even if members of smaller groups will always have more features in general in

common than the larger group that includes them That is they will have their

apomorphies their unique features as well as progressively more plesiomorphies

features found both in the small groups and in the larger clades of which they are a part

having more features in common is indeed one common meaning of more natural

(Note that the implication of the word natural has long been a group of the kind

[usually unspecified] that I think should be recognised and if other botanists had

different ideas too bad their groups were necessarily not natural [see Bather 1927] It

is thus rarely a helpful word and is not used here) Furthermore if this approach is

adopted we will find a slippery slope ahead By this kind of argument all families should

be very small since their members will have a great deal in common and so will be

most natural However as families (for example) are split the relationships that are

evident between the segregates and that were responsible for their being placed in a

single family in the first place will seem to necessitate the recognition of a new order

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

10

etc as is evident in Takhtajans own work - general taxonomic inflation is the result (see

also comparable suggestions in a cladistic context for Brassicales in particular - Ronse de

Craene amp Haston 2006) Such splitting is also questionable when teaching and learning

families since the student needs to understand the system as a whole However for

some genera removed from the families that until now have included them the

phenetic-classificatory-phylogenetic structure in their new home may mandate the

recognition of small families On the other hand Takhtajans suggestion that narrowly

defined families are more useful for phylogenetic studies may be true Indeed I have

more than once regretted prematurely combining groups whether species (in the

context of monographic work) or families (in the course of preparing these notes) but

this is largely a separate issue

Van Steenis (1978) Philipson (1987b) and others have rightly questioned the value

of splitting a group when ideas of the relationships of its constituent members have not

changed - that is very good reasons have to be provided for splitting a family if the

genera within it remain part of the same clade rather than belonging to another clade

Thus APG (2003) broadened the circumscription of Malvaceae because of the

parapolyphyly of some of the families that had historically been associated with it (Judd

amp Manchester 1997 Alverson et al 1999 Bayer et al 1999) These families particularly

Tiliaceae and Sterculiaceae were not at all easy to distinguish their close relationship

(see eg Brown 1814) and overall similarity had long been conceded and to some

workers at least their combination has come as something of a relief Although most of

the larger clades within Malvaceae sl remain difficult to distinguish even with flowers

Cheek (2007) opts for a wholesale and novel dismemberment into ten families

however the very good reasons for doing this are wanting

The same principles are of course applicable when it comes to dividing genera little

other than a headache is gained by splitting genera such

as Drosera and Gnetum (Doweld 2000) as has recently been proposed Thus if an

established genus divides into two (or more) clades this is not a signal for recognising

two groups at the same level - so here it could be argued that the dismemberment

of Pterostylis (Jones amp Clements 2002b) was somewhat unfortunate Along the same

lines if a newly-discovered taxon is sister to an existing named taxon - say a genus - this

does not necessitate the description of a separate genus for the newly described species

(cf Davis 2002) (Of course some [eg Thorne 1976] have suggested that the sizes of

gaps between groups at the same rank should be similar but any principle like this is

inherently flawed since morphological gaps are more unstable than phylogenetic

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

11

relationships and applying it across all flowering plants would both be difficult and cause

substantial changes - see also below) Humphreys and Linder (2009) provide a well-

documented survey of generic concepts in plants which the reader should consult they

note that generic limits (broad versus narrow) have oscillated historically and that

currently larger genera tend to be recognised because studies tend to be on a broader

scale than in the past

Note that invoking similarity or difference - whether qualified (considerable

similarities substantial differences) or not - in a cladistic context as justification for

combining or splitting taxa is not a particularly strong argument (see eg

CardiopteridaceaeStemonuraceae - Karingrehed 2002c) Similarity and difference can

neither be defined precisely since what may seem to be substantial similarities to me

may not to the next person nor are they likely to be stable in the face of our changing

knowledge of morphology and what might be synapomorphies

I might have prefered to merge some families recognised here or split others but

by and large I do not think my own preferences matter very much - and I take the same

position with regards to comparable preferences expressed by others Indeed the

bottom line is that in flagged hierarchies of the kind used here the limits of any

monophyletic unit generally taught and discussed particularly other than species can

be established only by convention and consensus (eg Stevens 2002 2006a Entwisle amp

Weston 2005) This is rather different from the reasonable observation that the

phylogeny itself cannot be achieved by consensus (Thorne 1976) for in most cases there

is a fact of the matter when it comes to relationships Given the increasing support for

the outlines of angiosperm phylogeny a stable consensus classification based on this

phylogeny seems attainable Indeed in addition to providing current ideas of

relationships of seed plants in a synthesised form this site is part of an attempt to build

such a consensus about the circumscription of taxa (see APG 1999 2003 2009 Grass

Phylogeny Working Group 2001 Mabberley 2008) Reaching such a consensus is vital

since what we know of angiosperm phylogeny allows a very large number of

classifications to be based on it and as we find out more the number of possible

classifications increases greatly Unfortunately however nature does not dictate what

the classification should be All classifications are constructed by humans to

communicate particular aspects of groups and relationships Our goals as systematists

are surely to produce robust hypotheses of relationships to understand the evolution of

morphology and the like - but not to argue ad nauseam whether something should be

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

12

a family or a subfamily That way surely lies madness and worse the discredit of our

discipline

There are similar issues whatever naming system is used Thus in phylogenetic

naming (Baum et al 1998 for an example but cf Baum et al 2004 for the PhyloCode

see Cantino amp De Queiroz 2006) an unflagged hierarchy is used in which any

terminations of names used are uninformative about the relative position of taxa If one

adopts the principle of phylogenetic naming one indeed does not have to worry about

the nomenclatural consequences caused by lumping or splitting any well-supported

clade can be named without affecting the name of more or less inclusive clades

Unfortunately unflagged hierarchies have very serious deficiences as communication

devices because they lack one aspect essential in language biological or otherwise -

they contain no intrinsic information about the relationships of the group in question to

others (eg Pfeil amp Crisp 2005 Stevens 2006a) Recent suggestions for using prefixes like

Apo- and Pan- to PhyloCode names will however allow limited information of this

kind to be conveyed but only as it pertains to individual branches and current

proposals do not even mandate that the prefixes be employed consistently In any

event such proposals simply prevent the potential tripling of the number of quite

different names used to describe different aspects of a phylogenetic tree over those

used to name monophyletic groups pure and simple In general where n is the number

of extant species in a group the number of clades in such a group = n-1 (Species will

also need names too for their names see Dayrat et al 2008) Importantly here too

consensus over the clade names commonly learned by students and used in herbaria is

needed otherwise communication will be impeded the names themselves will provide

no guidelines as to which should be chosen The situation is of course more complicated

than this Terminations that convey ideas of rank in a phylogenetic classification can also

be used in phylocode names - however there they will carry no implications of rank

How they will be used is another matter of course

Of course there are other philosophies of classification and some still prefer

evolutionary classifications There classificatory principles differ substantially from those

followed here eg the recognition of paraphyletic taxa may be permitted however

detailed reasons for prefering the taxa that are recognised are rarely given although

nature and natural groups are often mentioned (cf Stuessy amp Koumlnig 2008) For

summaries of commonly used systems see Brummitt (1992) and Mabberley (2008)

new evolutionary systems appear every year or so However even those who allow or

promote the recognition of paraphyletic groups (eg Grant 2003 Thorne 2007

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

13

Heywood et al 2007) may find it of interest to examine a system recognizing only

monophyletic groups it provides a rather different understanding of evolution

Evolutionary classifications in general try and combine phylogeny and

morphological gaps although that is no easy thing to do - it is akin to combining chalk

and cheese (for an attempt to make this impossible task seem to be more objective see

Stuessy amp Koumlnig 2008) However there is a principle from evolutionary classification

that is relevant and I think quite useful here The size of the gap between two groups

tends to be inversely proportional to the sizes of the groups involved (Davis amp Heywood

1963) One can imagine a situation in which a large group is formally divided even

although the distinguishing characters of the two are weak whereas a smaller group

similarly divisable will be left intact

To summarize If hypotheses of phylogeny remain stable we should be able to base

a stable classification on that phylogeny and then get on with our work that is testing

the phylogenies we have elucidating phylogenies in areas where relationships are

unclear studying the evolution of morphology describing species etc In this context

the spread of the Angiosperm Phylogeny Group system (see below) and its widespread

utilisation in technical literature also floras (eg van der Meijden 2005) textbooks (eg

Simpson 2006 Judd et al 2007 [third edition]) dictionaries (Mabberley 2008) more

popular literature (eg Souza amp Lorenzi 2005 Spears 2006) and as an outline for a new

herbarium sequence (Haston et al 2007) is gratifying The posibility that one might be

able to develop a stable phylogeny-based classification of families and in particular

orders represents a dramatic turn-around from the pessimistic attitude about such

higher-level groupings expressed by Davis and Heywood (1963) and Thorne (1976) the

latter even suggesting that we should bury forever the metaphor of the phylogenetic

tree as highly unrealistic (ibid p 56) Returning to Godfray and Knapps (2004) users

of classifications who want a stable informative and accessible classification that

enables easy identification - unfortunately they want cake with everything and cannot

get it - these pages attempt to satisfy as many of their needs as possible but phylogeny

and monophyly are the primary shapers of the APG classification

On this classification in particular

Here I very largely follow the Angiosperm Phylogeny Group classification (APG

2003) Any differences are not to be interpreted as differences in principle simply that

new phylogenies continue to be published and here I attempt to provide an overview

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

14

of current ideas of higher-level relationships of seed plants The Angiosperm Phylogeny

Group classification is based on well-supported relationships evident in the numerous

molecular studies that began to appear in the late 1980s (see APG 1999 for the

principles underlying the classification) but recent phylogenetic work does not

contradict the major outlines of the trees used by APG II (2003) or even those in APG

I (APG 1999) The APG classification has for the most part been conservative and I

am similarly conservative here For the names and authorities I follow APG III (2009)

although the names of the authors may not always be correct these being in something

of a state of flux for the superordinal names used see Chase and Reveal (2009)

although classification at this level here is not exhaustive - only clades with several

orders have superordinal names

Some changes in our ideas of relationships and hence in the clades we talk about

are particularly likely in parts of Lamiales Caryophyllales and Malpighiales for example

although this will not affect the orders themselves Thus some changes to clade

circumscriptions are to be expected although I do not expect them to be substantial

but changes are neither a defect of cladistics nor a necessary consequence of the use of

molecular data Clades are hypotheses of relationships and as hypotheses they may be

overturned However work since APG I has not suggested other than minor changes

in the compositions of the orders even if the odd genus or even family is turning out to

be seriously misplaced - recent examples are Hydatellaceae (from monocots-Poales to

Nymphaeales Saarela et al 2007) Guamatelaceae (from Rosales-Rosaceae to

Crossosomatales Oh amp Potter 2006) and Perrottetia and Bhesa (from Celastrales-

Celastraceae to Huerteales and Malpighiales respectively Zhang amp Simmons 2006) The

main changes have been clarification of the relationhips of individual families or groups

of families that were of uncertain position eg of Chloranthaceae (Moore et al 2007)

Ceratophyllaceae (Jansen et al 2007) and unplaced asterid II families (Winkworth et al

2008a) As mentioned above phylogenies in the area of main interest in these pages are

overwhelmingly tree-like Although there is evidence for acquisition of host plant genes

by parasites (see below) even wider but inexplicable transfer of mitochochondrial

genes in plants like Amborella (Bergthorsson et al 2004 cf Goremykin et al 2009a)

and even transfers of nuclear genes (Vallenback et al 2008) these are the exception

rather than the rule

In cases where the Angiosperm Phylogeny Group II allowed alternatives as to the

limits of families - Papaveraceae in the broad sense or Papaveraceae plus

Pteridophyllaceae plus Fumariaceae Proteaceae in the broad sense or Proteaceae plus

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

15

Platanaceae - the choices made here follow common usage eg as in textbooks like

Judd et al (2007) and Simpson (2006) and particularly in the new edition of

Mabberleys The Plant Book (Mabberley 2008) This latter is itself an attempt to reflect a

consensus the result of taking the opinions of botanists at several meetings A largely

similar consensus is reflected in the APG III classification (APG 2009) where you will

see that alternative classifications have been dispensed with For many people the

existence of such alternative classifications will simply confuse so agreement over

which groupings to use when alternatives are permitted and then dispensing with the

alternatives that are not commonly used seems reasonable I am making the few

changes that are needed to bring the classification here in line with these others but

only gradually since there is not enough time in each day

As already mentioned higher-level relationships in general and the composition of

orders in particular have always presented something of a challenge to systematists

(eg Davis amp Heywood 1963) That being said the composition of some clades like

Apiales Crossosomatales and Pandanales is decidedly unexpected However it is

interesting to see that these higher level clades are generally accepted even in works

with different classificatory philosophies (eg Heywood et al 2007) For clades like

Malpighiales many of the family groupings within Asparagales etc attempts to find

distinctive characters have largely failed (but see Endress amp Matthews 2006a also

the Apomorphies page here) Interestingly as with families some groupings suggested

by molecular studies are supported by morphological andor chemical characters that

have long been known sometimes for over a hundred years the relationship between

Pittosporaceae and ApiaceaeAraliaceae are a case in point (Hegnauer 1969b and

references) As our knowledge of morphology and chemistry improves we can hope for

improvements in the characterisations of clades at all levels

In many cases the new family limits of the Angiosperm Phylogeny Group (see

APG 1999 2003) are not really controversial although changes from the limits

commonly accepted only a decade ago are sometimes quite dramatic (eg Wagenitz

1997) Thus the split of the old Saxifragaceae sl is necessitated by its extreme

polyphyly as also with Icacinaceaesl and Cornaceae sl However the limits and

relationships of such groups had long been considered to be unsatisfactory but until

recently there had been no real reason to prefer one proposed arrangement over

another It is generally accepted that the limits of Lamiaceae and Verbenaceae have to

be redrawn although there is as yet no compelling evidence that the redrawn taxa are

not sister taxa (there is no evidence that they are) But whatever their relationships the

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

16

content of the clades has changed considerably and incidentally they are now easier to

identify than before the decision to recognise two families is not difficult The same is

true for Salicaceae and Achariaceae (Malpighiales) two previously small families that

have received the bulk of the old Flacourtiaceae Clade and hence taxon limits remain

difficult around Theaceae (Ericalessee alsoSladeniaceae Pentaphylacaceae)

and Euphorbiaceae (Malpighiales see

also Phyllanthaceae Picrodendraceae Putranjivaceae etc) Here current groupings

may not represent quite such dramatic changes in our understanding of relationships

Indeed parts of the old Euphorbiaceae that were separated may yet go back together

particularly Phyllanthaceae and Picrodendraceae (see Wurdack et al 2004 Davis et al

2005) even if Theaceae should indeed be dismembered (Geuten et al 2004) However

given that there is no molecular evidence that currently warrants combining all the

segregates of Euphorbiaceae and even if some do come together the clades the

families represent suggest novel groupings not recognised in current classifications

Furthermore Rafflesiaceae appear to be embedded within Euphorbiaceae s str (Davis

et al 2007) so the family is divided Of course maintaining even a moderately narrowly-

circumscribed Euphorbiaceae would entail reducing the iconic Rafflesiaceae to

synonymy Relationships in core Caryophyllales especially

around Phytolaccaceae andNyctaginaceae and also Portulacaceae are incompletely

understood and refashioning of taxon limits will doubtless be needed as cladistic

relationships become apparent (eg see Nyffeler 2007 Ogburn 2007a b) Some

groupings in the old Icacinaceaeand Olacaceae also remain unclear

The discovery of the relationships of parasitic and aquatic groups have presented a

particular challenge to systematists Morphologically some of these plants are so highly

modified that interpretation of the plant body in conventional terms is difficult or even

impossible Thus parasitic groups (broad-leaved angiosperms only) such as Rafflesiaceae

are hard to place since both the vegetative body and the flowers are changed almost

beyond recognition (flowers of rafflesiaceae are a good example of gigantism - Davis et

al 2007) Furthermore plastid gene sequences may be difficult or impossible to obtain

the chloroplast DNA in particular being highly degraded and the rate of molecular

change in general may be high so the problem of long-branch attraction is serious (eg

Duff amp Nickrent 1997 Nickrent et al 1998 Caddick et al 2002a G Petersen et al

2006b) Echlorophyllous sapromycoheterotrophic taxa mostly known from monocots

present similar problems However progress is being made placements for

Apodanthaceae Rafflesiaceae Mitrastemonaceae Cytinaceae and Cynomoriaceae have

recently been suggested (Barkman et al 2004 2007 Davis amp Wurdack 2004 Nickrent et

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

17

al 2004 Davis et al 2007) it seems likely that Burmanniaceae sl are polyphyletic but

both parts are in Dioscoreales (Merckx et al 2006) and relationships within the largely

hemiparasitic Santalales are also gradually being clarified (Maleacutecot 2002) The inclusion

of parasitic taxa in general molecular analyses can cause conniptions (eg Nickrent et al

2004 Davis et al 2004 Chase et al 2006 G Petersen et al 2006b) and there can be

horizontal transmission of genes (eg Davis amp Wurdack 2005 Vitaceae to Rafflesiaceae

Barkman et al 2007 the mitochondrial atp1 gene commonly moves)

Similarly in water plants neither vessels in particular nor much xylem in general is

needed leaves are highly modified and water-mediated pollination if adopted may

well be associated with major changes in floral morphology Here too recent molecular

studies suggest that aquatic groups with hitherto problematic relationships may find

homes Thus Podostemaceae are close to Clusiaceae and Calophyllaceae and are sister

to Hypericaceae (Malpighiales Kita amp Kato 2001) Hydatellaceae which used to be in

Poales are part of Nymphaeales (Saarela et al 2007) and Hydrostachyaceae may be

close to Hydrangeaceae (Cornales Xiang et al 2002) or perhaps should be placed in

Lamiales note that in the first two cases in particular there are morphological and

chemical features that support such a move If Podostemaceae are indeed close to

Clusiaceae I look forward to seeing hypotheses to explain how the dramatic changes in

the vegetative body that have made Podostemaceae so problematic for generations of

systematists took place That conventional wisdom has trouble in understanding or

explaining how the morphologies of groups like Clusiaceae and Podostemaceae can be

related is largely a problem with conventional wisdom Ceratophyllaceae seem to be

finding a position as sister to eudicots (see Moore et al 2007) but their morphology is

so derived that there is no morphological evidence of which I am aware for this

relationship There are also similar although less extreme morphological problems with

many plants pollinated by wind

In such situations especially with some water plants mycoheterotrophs and

parasites a variety of characters may be affected and particularly under such

circumstances groups have tended to be formed using a few or even only a single

character that seems to provide evidence of relationships Add to this the tendency to

weight some characters particularly strongly on a priori grounds and the result was

either the recognition of conglomerate taxa such as Amentiferae which now appear to

be highly polyphyletic or the segregation of families like Plantaginaceae s str (now

much expanded) and Leitneriaceae (now in Simaroubaceae) - all of these are more or

less wind pollinated

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

18

Even although relationships of many parasitic and aquatic groups are being

clarified and at least some of the proposed phylogenetic positions seem to be well

supported it is indeed often still very difficult to understand how such groups have

evolved from their more morphologically conventional relatives However we have had

all sorts of preconceptions as to how characters should evolve Thus we used to

assume that features like highly scalariform vessel perforation plates or the complete

absence of vessels or a flower with an androecium that had many stamens a superior

ovary or separate petals were necessarily plesiomorphic or primitive and conversely

a simple perforation plate and the presence of vessels an androecium with few

stamens inferior ovary or petals that were connnate were almost necessarily

apomorphic or advanced (but cf eg Stebbins 1951) Such assumptions are incorrect

(eg Soltis et al 2005b) Carpels may become secondarily free carpels may fail to close

the seeds then developing outside the confines of the carpel as in some Aspagaraceae-

Nolinoideae Violaceae Berberidaceae Malvaceae-Sterculioideae etc) in Peliosanthes

teta perhaps the only species in Peliosanthes(Asparagaceae-Nolinoideae) the ovary

varies from superior to inferior (Jessop 1976 species limits here need close

investigation see also Kuzoff et al 2001 and Soltis amp Hufford 2002 Saxifragaceae

Apiales etc) many-seeded carpels can evolve from few-seeded carpels

(Razafimandimbison et al 2008) monoecy may be derived from dioecy (Schaefer amp

Renner 2010 and references) Classic studies such as those by Babcock (eg 1947)

on Crepis that assumed that evolution - in this case of the karyotype in particular - was

unidirectional have needed comprehensive re-evaluation (Enke amp Gemeinholzer 2008)

Most if not all characters have reversed andor evolved in parallel as is clear in the

discussion of ovary position here especially in Poales Saxifragales and Asparagales and

also in comments on such morphologically distinctive taxa (when compared with their

immediate relatives) such as Menyanthaceae and especially Pittosporaceae Parallel

evolution may occur even at the level of amino acid substitution as in the independent

acquisition of the phosphoenolpyruvate carboxylase (pepC) gene in C4 photosynthesis in

grasses (Christin et al 2007b see also Blaumlsing et al 2000)

Thinking of how such characters have been used in the past in classifications at the

generic level the results have been similar to those just discussed above for parasitic

plants and others Relying too much on animal pollination syndromes to mark generic

boundaries has all too often led to taxa that are highly unsatisfactory phylogenetically

(see eg Acanthaceae Bignoniaceae Campanulaceae Ericaceae Lamiaceae

Orchidaceae) and over-reliance on characters of fruit and seed (see eg Brassicaceae

and Apiaceae) has also led to unsatisfactory generic limits Again the more general

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

19

problem is the use of one or a very few characters that have been weighted a priori to

structure classifications (see also Garciacutea et al 2009) although it is perhaps not

surprising that such an approach sometimes results in the delineation of taxa that have

indeed turned out to be quite distinct (cf eg Burtt Astragalus versus Oxytropis)

Taxa recognized here are monophyletic yet many are polythetic at the

morphological level that is they lack unique features characterizing (defining) all and

only members of that family they can be recognised phenetically only by the unique

combinations of characters that they posess This is the result of evolution any

synapomorphy characterizing a taxon may be lost or modified beyond easy recognition

in some of its members or the synapomorphy may appear to be identical to a feature

that has evolved in parallel in a quite unrelated plant That plant groups are polythetic is

almost as much a feature of monophyletic taxa as the paraphyletic taxa common in

evolutionary classifications Some families now include substantial variation as

phenetically distinct derived groups are placed in their proper phylogenetic position -

examples are Ericaceae which include the erstwhile Empetraceae a wind-pollinated

group that is florally very different and the overwhelmingly large-flowered and animal-

pollinated Plantaginaceae which now include several derived small-flowered aquatic

and wind-pollinated groups that were previously placed in separate families

Thus some families as delimited here may not be easy to recognize However

remember that detecting relationships - use whatever characters you can even if they

are not obvious - and naming a plant - focus on easy-to-see characters that may not

reflect relationships - are quite different problems Taxa although natural may not be

readily recognizable indeed it was in exactly this context that Lamarck worked out the

basic principles of writing dichotomous keys in 1778 (see Scharf 2007 for keys and the

like) Of course Lamarcks idea of nature was very different from ours - he thought that

there was some kind of continuum of form on which living organisms were to be

situated with no real gaps anywhere - but this meant that his genera (for example)

might well not be sharply distinct from each other even if each were part of the real

continuum that was life Lamarcks separation of the establishment of relationships and

groupings in a system and identification of the plants in that system have been widely

accepted (but cf eg Godfray amp Knapp 2004 many of the contributors to Heywood et

al 2007) Perhaps the best way of identifying plants at the family level is by well-made

multiple access keys as in Watson and Dallwitz (1992a onwards family limits there may

differ substantially from those adopted here) Multiple access keys free users from the

constraints of dichotomous keys in which particular characters are needed at each step

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

20

of the identification process before there can be further progress Instead those

characters that happen to be evident on a specimen can be used in whatever order is

convenient when linked to illustrations glossaries etc their power is enormous (see

Dallwitz et al 2000 [2006] for the principles underlying their construction and use)

Nevertheless dichotomous keys such as those of Hutchinson (1973) and Franz Thonner

(Geesink et al 1981) have their uses Of course taxa coming out adjacent in keys may

well not be at all related

(When identifying large numbers of plants even more efficient than either style of

identification and certainly lots more fun is sight identification Unless you have a

photographic memory you have to build up your knowledge of comparative plant

morphology - on which the ability to make accurate identifications depends - by

repeated observation When faced with an unknown plant I always look for leaf teeth

and stipules distinctive hair types especially stellate and T-shaped hairs glands and

punctations of any sort the presence of latex or other exudate and check leaf insertion

smelling crushed leaves can also be helpful In this context nodal anatomy can usually

be checked using a razor and a hand lens or even simply carefully examining leaf scars

The short paragraphs added after most families may help in confirming familial

identifications)

For the record and for the little that it is worth there are 4 orders and 13 families

of gymnosperms characterised on these pages and together they include some 82

genera and 947 species For angiosperms comparable figures are 56 orders 445

families 13208 genera and 261750 species (of which monocots include 11 orders 89

families 2759 genera and 52760 species) Note however that higher mathematics

was never my strong point and anyway these are pretty meaningless figures even for

species which many (but not all) might concede smacked slightly more of reality than

other taxa estimates range as high as 422000 (Govaerts 2001) Furthermore numbers

of genera and species change daily Nevertheless as emphasized here families are

useful in teaching we as a community can ensure that their limits remain largely stable

and by concentrating on relatively few of them one can gain some familiarity with much

of the worlds flora For summaries of the sizes of orders and families and of the general

arrangement followed in these pages see also the Statistics page itself

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

21

A PHYLOGENETIC TREE OF SEED PLANTS (THE MAIN TREE)

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

22

The Main Tree above is a conservative summary of well supported relationships

between the major clades of extant seed plants Reading - interpreting and

understanding - such a phylogenetic tree is not necessarily easy so reading a tree may

require a bit of practice OHara (eg 1988 1992) provides an accessible introduction to

thinking and talking about history in the context of looking at phylogenetic trees More

recently Krell and Cranston (2004) Crisp and Cook (2005) and others have emphasized

how careful one must be when interpreting and talking about evolution in ladderized

trees in particular and phylogenetic trees in general Gregory (2008) provides a

particularly detailed discussion of this problem In ladderized trees the smaller (in terms

of numbers of terminals) sister taxon is consistently shown on the same side at every

node thus the trees tend to be pectinate like the teeth of a comb however the

horizontal axis that is as a result so evident has no polarity or particular direction The

use of the adjective basal when discussing phylogenetic trees is especially dangerous

(see also Wojciechowski et al 2004 D Soltis et al 2005b) When I use the term and the

context is not otherwise clear I am referring to the pectinations at the base of a

ladderised tree Remember that when talking about sister taxa one can never be basal

to the other or older or younger although members of one can be more derived - in the

sense of having more apomorphies - than the other (but even then one has to be

careful) The words primitive and advanced and lower and higher should

practically never be used especially when talking about taxa Amborellaceae and

Pinaceae are sister to all other flowering plants and Pinales and are basal to the crown

groups of all other flowering plantsPinales respectively but that does not mean they

are thereby primitive Pinaceae in particular have numerous apomorphies The word

plesiomorphic is far less loaded than primitive and can be used to talk about

individual characters

Most trees here have been more or less ladderized as is the one above The

ladderization may be imperfect for example you can see that on the Main Tree

asterids with ten orders follow rosids and relatives which have seventeen although

this is in part because of the currently unresolved nature of relationships between the

main clades of core eudicots (this is likely to change soon) The Main Tree could have

been drawn with Amborella Acorus or a host of other taxa at the far right without

offending any relationships phylogenetic trees are like mobiles the only fixed points

being the nodes Despite the problems just mentioned pectination interpreted

carefully has its value and it is not an arbitrary process As one reads the terminals of a

pectinate tree from left to right adjacent terminals will be separated by apomorphies

that will cumulate in an appropriate fashion When thinking of a book or a herbarium

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

23

sequence (see Haston et al 2007) this is of some value Nymphaeales and

Austrobaileyales are here adjacent on the tree but they could be separated by

hundreds of families in the sequence merely by rotating only the node from which

Nymphaeales and their sister group (all other angiosperms minus about 75 species)

arise If adjacent in a book or herbarium or as in these summary pages then it is

relatively easy to relate their characters both apomorphic and plesiomorphic but if

separated by hundreds of pages or two floors in a large building then it is less easy to

get anything from the sequence (Since all orders in the pages of the main part of this

site are preceded by the apomorphies of all nodes immediately below them in the seed

plant phylogeny and because of the linkages that have been built in to the site this

problem is much less serious there there is no sequence) So the particular sequence of

all seed plants that is based on a phylogenetic classification but is to be used in herbaria

or any other place where taxa have to be arranged linearly as in this part of APweb

will be one that maximizes the number of taxa that are both successive branches of the

tree and placed successively in the sequence Since specimens are generally filed under

families the outline of a new family sequence for arranging herbaria and books can now

be suggested (Haston et al 2007) and such a standardized sequence will further help

teaching and learning about plants

ON THINKING ABOUT APOMORPHIES

Identifying apomorphies is important because understanding the evolution of

morphology in the broad sense - ie including anatomy chemistry etc - in the context

of diversification is one of our major goals For identifying apomorphies several

preconditions must be met One needs to have an accurate well supported phylogeny

one has to have examined the right taxa both from the point of view of morphology and

molecules one has to have coded the characters right (ie delimited states

appropriately) and one has to use an appropriate model of evolution when fitting the

variation to the tree (see eg Omland 1999 Stevens 2006b) If the distributions of

apomorphies below are compared with those in other studies (eg Turgeon et al 2001

Bremer et al 2001 Endress 2001 Albach et al 2001a Judd amp Olmstead 2004 D Soltis

et al 2005b Zhang et al 2006 Judd et al 2007) the reader may find a number of

differences Although I have integrated such studies as far as is possible there are five

reasons that link to the preconditions just mentioned why there may be differences

where features are placed on trees

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

24

1 I may not have found all the information about a particular character there may be

disagreement over its interpretation or I have added information

2 The sampling of nearly all molecular studies is very incomplete (see Salisbury amp Kim

2001 for problems caused by sampling) indeed it is commonplace to decry the

incompleteness of molecular sampling But not only is the sampling in molecular studies

often less than we might wish that of the morphological and chemical characters whose

evolution we are interested in understanding is also often very poor So for many

anatomical chemical and embryological characters that are confidently said to

characterise families and other groups we all too often have no idea if those characters

are applicable to the whole clade or just to a subgroup within that clade However as

our sampling improves we can locate changes on the tree more precisely if in

unexpected positions Thus is is sometimes suggested that Ericaceae have ellagic acid

(Soltis et al 2005b) in fact the little evidence we have indicates that only a small clade

of some 80 species has ellagic acid and the rest of the family - itself alone about one

third of all Ericales - does not Similarly Ericaceae are noted for their

ectendomycorrhizal associations but Enkianthus sister to all other Ericaceae

apparently lacks these (Abe 2005) Thus these mycorrhizae and their associated

distinctive hair roots are unlikely to be apomorphies of Ericaceae - however it appears

that other families near Ericaceae may also have ectendomycorrhizae (see Asai 1934)

3 I am fitting characters to a very conservative tree with many polytomies although the

nodes that are utilised are for the most part strongly supported polytomies make the

optimisation of characters that is the assigment of character state change to a

particular node on the tree notably difficult (eg Madison amp Madison 2002) In nearly

all studies of the evolution of characters distributions of characters are optimised on a

more or less fully resolved tree and the construction of supertrees may yield yet more

detailed hypotheses of relationships (for literature on supertrees see Cotton amp

Wilkinson 2007 2008) Of course some nodes on such fully resolved trees andor

supertrees may have little support and optimisations of characters on such trees may

carry correspondingly little conviction Even parts of some of the trees used here have

poor support eg relationships within aquatic Alismatales etc although I indicate

when this is the case

4 Exactly how one goes about optimising a character on a tree is critically important

Even using simple parsimony optimisations (ACCTRAN or DELTRAN ACCelerated

TRANsitions or DELayed TRANsitions) the position of synapomorphies on trees - and

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

25

hence our ideas of evolution - may differ and this is still more true if one uses maximum

likelihood or Bayesian methods and makes apparently reasonable suggestions about

weighting gains over losses (eg Donoghue amp Ackerley 1996 Cunningham et al 1998

Omland 1997 1999 Ree amp Donoghue 1999 Polly 2001 Webster amp Purvis 2001

Ronquist 2004 Crisp amp Cook 2005) Sannier et al (2007) show how in in Arecaceae that

where on a phylogenetic tree one might peg changes in microsporogenesis will depend

on the methods one uses to do this and Pedersen et al (2007) discuss the sometimes

very substantial effect of node support on the posterior probabilities of ancestral

character states Here I use parsimony optimization not always as explicit as it might be

but I have often indicated where there are particularly important uncertainties as to the

positions of particular character changes on the tree

5 Finally although I have paid quite a lot of attention to the delimitation of the

character states that make up all the characterizations I have not spent enough time on

this critical operation If we are interested in understanding evolution then fitting the

basic variation - not character states - to a tree in principle allows greater flexibility in

understanding morphology in the context of local phylogenies (see also Stevens 2000

Endress 2005c) However many character states used here are delimited globally that

is they are circumscribed in the context of the variation shown by individual characters

across all angiosperms andor in the context of classic ideas of character evolution

Character states often have arbitrary limits and serve best to communicate

information whether they are in fact suitable for either phylogenetic analysis or

understanding evolution are separate issues Studies have rather unsurprisingly

perhaps but importantly shown that dividing the one character into different sets of

states may yield differing ideas of evolution of that character (eg Lamb Frye amp Kron

2003 Hibbett 2004) When looking at trees on which character states are optimised

one should bear in mind the problems surrounding the delimitation of states (eg

Stevens 2000 2006b) and the danger of using pollen or other types - constructs

based on many characters that vary independently but which effectively get lost in

these types thus Blackmore et al (2009) decomposed the pollen types in Asteraceae

into 52 characters

I use Remanes three main criteria of homology or better similarity when

determining the basic similarity of structures on different organisms (see Remane 1952)

These criteria are special properties position and intermediates Special properties

include anatomical or chemical characters gene expression data etc Position refers

to the position of an organ with respect to landmarks on the plant Although plants are

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

26

plants and landmarks are not as fixed as one might like trying to work out the relative

positions of parts is a good way of understanding morphology so long as one does not

try to out-do Procrustes Intermediates include intermediates found on the same

plant or on different plants Intermediates may observed during development or be

adult structures Thus one may find changes during development which suggest what

the nature of a particular structure is and very different stuctures on different plants

may turn out to be similar early in development Or one can simply compare different

taxa and see that structures that appear to be very different when comparing two

different taxa can be linked morphologically by looking at other taxa

Nevertheless the use of these criteria may not yield an unambiguous answer as to

what a structure is even given a solid phylogeny and an improved understanding of

development (see Jaramillo amp Kramer 2007 for a useful discussion) As Endress (2005c)

observed a number of features - position function development shape anatomy

histology gene activity and relationships to other taxa that clearly have petals - can be

used to distinguish a petal (for example) from other floral structures if a petal does not

have one of these features is it thereby not a petal Thus Maturen et al (2005) recently

found that floral organ diversity genes (B and C) were expressed in the large white

inflorescence bracts of Cornus (see also Costa et al 2005) Peney et al (2005) noted that

not all monosulcate pollen grains in monocots have the same developmental pathway

and that as a result such pollen might not have the same ancestral state Reeves and

Olmstead (2003) suggested that the genetic mechanisms causing monosymmetry in

Lamiales and Solanales were different and Serna and Martin (2006) described similar

problems with the development of hairs in Arabidopsis when compared with that of

hairs in Antirrhinum and Solanaceae Indeed as one perhaps might expect delimitation

of states and characters does not necessarily become easier with increasing knowledge

of development etc Thus Buzgo et al (2004) Matthews and Endress (2005) and others

have shown how hard it can be to distinguish between eg prophylls and other floral

structures as their behaviour is studied during the course of the development of the

flower Similarly at what concentration is a particular secondary metabolite deemed to

be present (Waterman (2007)

The validity of the approach used here that of fitting morphological variation to a

largely molecular-based tree may be questioned However I think it rather unlikely that

well-supported molecular branches will be overturned by morphological data Indeed

analyses of morphological data alone do provide support for many of the clades evident

in molecule-only analyses and analyses with morphological and molecular data

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

27

together may lead to increased support for clades (eg Hufford 1992 Nandi et al 1998

[but adding morphological data reduces support for a number of critical clades too]

Doyle amp Endress 2000 however in none of these papers is the use of morphology

without ambiguity) It is unfortunately clear that the use of morphology alone may not

suggest problems in the phylogenetic placement of taxa that later turn out to have been

wrongly included (eg Zhang et al 1992) Thus I have been wary of putting much weight

on clades that have only morphological support but note that I have treated molecule-

based clades with low bootstrap or jacknife support values (esp below 70) or low

posterior probabilies (below 095) likewise Although I may have been mistaken in

placing so much emphasis on molecular data in terms of providing the basic

phylogenetic framework for angiosperms morphological and molecular data are only

very rarely in irreconcilably strong conflict There are indeed a few places where the

conflict seems extreme These include the relative positions of the Monimiaceae and

Hernandiaceae (Laurales) the position of Hanguanaceae (Commelinales [as here] or

Zingiberales) and of Triplostegia (is it in Dipsacaceae or Valerianaceae - see

Dipsacales) Fossils are unlikely to affect the topologies of the trees presented here but

see below for their importance in understanding morphological evolution in general and

the evolution of angiosperms in particular Nevertheless some largely reject the idea

that trees based on molecular data alone can recover phylogenetic relationships

especially when branching points are old and prefer to used trees based on analysis of

morphological data including those taken from fossils (Hilton amp Bateman 2006 Farjon

2007)

All in all however the extent of the congruence between morphological and

molecular data is impressive and heartening and many clades can be characterised

morphologically It seemed in 1998 that there were no unambiguous morphological

synapomorphies for angiosperm orders (K Bremer 2000) and this is still true if by

unambiguous is meant non-homoplasious However many orders and other clades

have synapomorphies even if these may be indistinguishable at least at the current

level of morphological and developmental knowledge from parallel occurences

elsewhere As our morphological knowledge increases so too does the number of

apomorphies

Endress and Matthews (2006a) emphasize the importance of tendencies and

developmental constraints when thinking about characters of clades within the rosids -

many characters or character combinations occur in a rather sporadic fashion within

the clade and apparently notably less frequently outside it There are a number of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

28

examples of tendencies here and in the individual order pages such as the features

enclosed by parentheses in groups above the level of family eg see (cuticular waxes

as aggregated rodlets) for the commelinids and the discussion of the distribution of

polyandry that is flowers with numerous stamens in the asterid I + II groups etc

Tendencies may involve more than single characters When thinking of the

associations of features like integument thickness and vascularization of the integument

(Endress amp Matthews 2006a) one can imagine fairly simple developmental

preconditions being operative As Endress and Matthews (2006a) note it is difficult to

imagine vascular tissue developing in an integument only two cells thick Changes here

would be loosely correlated if morphologically linked However the strongly correlated

changes noted by Givnish et al (2005) are ecologically linked but are presumably

morphologicallydevelopmentally independent When there seem to be characters

evolving more or less together tests can be carried out to see if the changes are

concentrated on certain branches of the tree (eg Maddison 1990 Sanderson 1991

Maddison amp Maddison 2000) Such tests have rarely been carried out at higher levels in

angiosperms

One of the most striking examples of a tendency is the distribution of N-fixation

restricted as it is to a monophyletic group of four clades although it has arisen seven (or

perhaps several more) times independently within the N-fixing clade and several

members of two quite different kinds of bacteria are involved (eg Clawson et al 2004

Elliott et al 2007 Sprent amp James 2007) Other examples of work that bears on the issue

of tendencies include the findings that flowers of polysymmetrical Arabidopsis have

genes like TCP1 that are expressed asymmetrically during early development and TCP1

is a probable orthologue of the well-known CYC gene of Antirrhinum that is involved in

the development of monosymmetric slowers there (Cubas et al 2001 Costa et al 2005

etc) Parallelism might build on this underlying morphologically cryptic monosymmetry

even if details of the genetic mechanisms causing the monosymmetry evident in

particular groups may be different (Reeves amp Olmstead 2003 Cubas 2004 see above)

Similarly there have been several recent suggestions that the capability to synthesise a

particular metabolite may be switched off but not lost and so can sometimes be

reacquired (eg Wink amp Witte 1983 Wink 2003 Liscombe et al 2005 Larsson 2007

Waterman 2007) Hence perhaps the rather spotty distribution of many secondary

metabolites like ellagic acid the indole alkaloid camptothecin iridoids etc when

considered in the context of phylogenies As a non-botanical example - but a rather nice

one - Salwini-Plawen and Mayr (1961) suggested some time ago that there has been

considerable parallelism (40-65 or more independent origins) in the evolution of eyes in

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

29

metazoans However Pax 6 seems to be a master control gene that is involved in eye

formation perhaps throughout all bilateralians (eg Gehring amp Ikeo 1999 Erwin amp

Davidson 2002) which changes the nature of the problem Note however that in

general our invocation of evolutionary tendencies to explain the patterns of

distributions of characters is really an admission that we do not understand those

patterns

As we find out more about variation we will probably find fewer and fewer features

constant throughout a group Most unqualified statements of presence and absence

should properly be qualified as usually present or usually absent if one is thinking of

the characterisations on the order pages as encompassing the total variation within a

clade Thus Pistia alone among monocots as so far known has sieve tube plastids with

starch grains not protein crystals However this does not affect the fact that sieve tube

plastids with cuneate protein crystals and lacking starch grains are an apomorphy for

monocots Furthermore fossils in a number of cases suggest character combinations

unknown in extant taxa as may be seen in the discussions of Fagaceae Platanaceae

Iteaceae Calycanthaceae etc Confusing the issue there may be questions as to where

exactly on the tree a particular fossil is to be placed (eg see Nymphaeaceae

Calycanthaceae Archaefructus etc)

To summarize given our current understandings of both phylogenies and

characters evolution of some characters in which we are interested seems very labile

(see eg D Soltis et al 2005b Endress and Matthews 2006a Stevens 2006b Ekman et

al 2008 for a good recent study) and I have been cautious when talking about character

evolution Much effort must continue to be spent in summarizing characters of well-

established clades at all levels providing features by which they may be recognized and

signaling synapomorphies Remember that (1) the basic morphological anatomical and

chemical knowledge of many critical taxa is woefully incomplete (2) different

assumptions about character evolution may greatly affect the position of

synapomorphies on trees (3) in many cases relationships within and between many

groups are too uncertain at present to worry very much about synapomorphies and

(4) we must be clear about what we do and do not not know As mentioned above I

have indicated in a number of places where there are particularly important

uncertainties as to where characters should be placed on the tree Nevertheless it is a

relatively easy matter to update notes such as these and it can be a simple matter to

incorporate new data on characters that have never before been considered in the

context of a tree Much basic - and unfortunately perhaps unfashionable - work must be

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

30

carried to clarify the distribution of morphological anatomical and chemical characters

for examples of what can to be done see the work of P K Endress and collaborators

(floral morphology and development) and S R Jensen and collaborators (iridoids) But

acquiring information about nodal anatomy cell and tissue distribution nodal anatomy

and the like is not difficult at all and should be generally encouraged

But all this is merely a necesary prelude to the understanding of evolution For this

one needs to know a lot more including dating the phylogeny understanding the

function(s) of characters factoring in relevant aspects of the palaeoenvironment in

which an apomorphy first appeared etc

SUMMARY OF THE SYSTEM

Below is a formalised summary of the relationships within orders of the families of seed

plants There are a few families that are not recognised even as options in APG II and

vice versa as well as a few extra orders The families to be recognised in the most

recent edition of Mabberleys The Plant Book (Mabberley 2008) are also largely

consistent with those below But all differences are trivial and will - I hope - eventually

disappear

Square brackets - [] - enclose clades the plus sign - + - designates sister taxa a

comma - - denotes part of a polytomy and quotation marks - - denotes a

paraphyletic group

SEED PLANTS

GYMNOSPERMS

Cycadales

Cycadaceae + Zamiaceae

Ginkgoales

Ginkgoaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

31

Pinales

Pinaceae [[Araucariaceae [Podocarpaceae + Phyllocladaceae]] [Sciadopityaceae

[Taxaceae + Cupressaceae]]]

Gnetales

Ephedraceae [Gnetaceae + Welwitschiaceae]

ANGIOSPERMSFLOWERING PLANTS

Amborellales

Amborellaceae

Nymphaeales

Hydatellaceae [Cabombaceae + Nymphaeaceae]

Austrobaileyales

Austrobaileyaceae [Schisandraceae + Trimeniaceae]

Chloranthales

Chloranthaceae

MAGNOLIIDS

Magnoliales

Myristicaceae [Magnoliaceae [[Himantandraceae + Degeneriaceae] [Eupomatiaceae +

Annonaceae]]]

Laurales

Calycanthaceae [[Siparunaceae [Gomortegaceae + Atherospermataceae]] [Monimiaceae

[Hernandiaceae + Lauraceae]]]

Canellales

Canellaceae + Winteraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

32

Piperales

[Hydnoraceae + Aristolochiaceae] [Piperaceae + Saururaceae]

MONOCOTS

Acorales

Acoraceae

Alismatales

Araceae [Tofieldiaceae [[Alismataceae [Hydrocharitaceae + Butomaceae]]

[Scheuchzeriaceae [Aponogetonaceae [Juncaginaceae [Maundiaceae [[Posidoniaceae

[Ruppiaceae + Cymodoceaceae]] [Zosteraceae + Potamogetonaceae]]]]]]]

Petrosaviales

Petrosaviaceae

Dioscoreales

Nartheciaceae [[Taccaceae + Thismiaceae] [Burmanniaceae + Dioscoreaceae]]

Pandanales

Velloziaceae Triuridaceae Stemonaceae [Pandanaceae + Cyclanthaceae]

Liliales

Corsiaceae [Campynemataceae [Petermanniaceae [Colchicaceae + Alstroemeriaceae]]

Melanthiaceae [[Philesiaceae + Rhipogonaceae] [Smilacaceae + Liliaceae]]]

Asparagales

Orchidaceae [[Boryaceae [Blandfordiaceae [Lanariaceae [Asteliaceae + Hypoxidaceae]]]]

[[Ixioliriaceae + Tecophilaeaceae] [Doryanthaceae [Iridaceae [Xeronemataceae

[Xanthorrhoeaceae [Alliaceae + Asparagaceae]]]]]]]

COMMELINIDS

Unplaced

Dasypogonaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

33

Arecales

Arecaceae

Poales

[Typhaceae + Bromeliaceae] [Rapateaceae [[Thurniaceae [Juncaceae + Cyperaceae]]

[[Mayacaceae [Eriocaulaceae + Xyridaceae]] [Flagellariaceae [Anarthriaceae

[Centrolepidaceae + Restionaceae]]] [Joinvilleaceae [Ecdeiocoleaceae + Poaceae]]]]]]]

Commelinales

[Commelinaceae + Hanguanaceae] [Philydraceae [Haemodoraceae + Pontederiaceae]]

Zingiberales

Musaceae [Strelitziaceae + Lowiaceae] Heliconiaceae [[Cannaceae + Marantaceae]

[Costaceae + Zingiberaceae]]

Ceratophyllales

Ceratophyllaceae

EUDICOTS

Ranunculales

Eupteleaceae [Papaveraceae [[[Lardizabalaceae + Circaeasteraceae] [Menispermaceae

[Berberidaceae + Ranunculaceae]]]]

Sabiales

Sabiaceae

Proteales

Nelumbonaceae [Platanaceae + Proteaceae]

Trochodendrales

Trochodendraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

34

Buxales

Haptanthaceae [Buxaceae + Didymelaceae]

CORE EUDICOTS

Gunnerales

Gunneraceae + Myrothamnaceae

Dilleniales

Dilleniaceae

Saxifragales

Peridiscaceae [[Paeoniaceae [Altingiaceae [Hamamelidaceae [Cercidiphyllaceae +

Daphniphyllaceae]]]] [[Crassulaceae [Aphanopetalaceae [Tetracarpaeaceae

[Penthoraceae + Haloragaceae]]]] [Iteaceae [Grossulariaceae + Saxifragaceae]]]]

Cynomoriaceae unplaced

Vitales

Vitaceae

ROSIDS

FABIDROSID I

Zygophyllales

Krameriaceae + Zygophyllaceae

Celastrales

Lepidobotryaceae + Celastraceae

Oxalidales

Huaceae [[Connaraceae + Oxalidaceae] [Cunoniaceae [Elaeocarpaceae [Brunelliaceae +

Cephalotaceae]]]]

Malpighiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

35

[Achariaceae [Goupiaceae [Violaceae + Passifloraceae] [Lacistemataceae Salicaceae]]

[Lophopyxidaceae Putranjivaceae] [Ctenolophonaceae [Erythroxylaceae +

Rhizophoraceae]] Linaceae Ixonanthaceae Humiriaceae Irvingiaceae

Centroplacaceae Pandaceae [[Bonnetiaceae + Clusiaceae] [Calophyllaceae

[Hypericaceae + Podostemaceae]]] [Malpighiaceae + Elatinaceae] Ochnaceae

[Peraceae [Rafflesiaceae + Euphorbiaceae]] [Phyllanthaceae + Picrodendraceae]

[Balanopaceae [[Trigoniaceae + Dichapetalaceae] [Chrysobalanaceae +

Euphroniaceae]]] Caryocaraceae

N-FIXING CLADE

Fabales

Quillajaceae [Fabaceae [Polygalaceae + Surianaceae]]

Rosales

Rosaceae [[Barbeyaceae [Elaeagnaceae Dirachmaceae Rhamnaceae]] [Ulmaceae

[Cannabaceae [Moraceae + Urticaceae]]]]

Cucurbitales

Anisophylleaceae [[Corynocarpaceae + Coriariaceae] [Cucurbitaceae [Tetramelaceae

[Datiscaceae + Begoniaceae]]]] Apodanthaceae

Fagales

Nothofagaceae [Fagaceae [[Myricaceae + Juglandaceae] [Casuarinaceae

[Ticodendraceae + Betulaceae]]]]

MALVIDROSID II

pgtGeraniales

Geraniaceae [[Melianthaceae + Francoaceae] [Vivianaceae + Ledocarpaceae]]

Myrtales

Combretaceae [[Onagraceae + Lythraceae] [[Vochysiaceae + Myrtaceae]

[Melastomataceae [Crypteroniaceae [Alzateaceae + Penaeaceae]]]]]

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

36

Crossosomatales

[Staphyleaceae [Guamatelaceae [Crossosomataceae + Stachyuraceae]]] [Aphloiaceae

[Geissolomataceae + Strasburgeriaceae]]

Picramniales

Picramniaceae

Sapindales

Biebersteiniaceae Nitrariaceae [Kirkiaceae [Anacardiaceae + Burseraceae]]

[Sapindaceae [Simaroubaceae Rutaceae Meliaceae]]

Huerteales

Gerradinaceae [Dipentodontaceae + Tapisciaceae]

Malvales

Neuradaceae [Thymelaeaceae [Sphaerosepalaceae Bixaceae [Cistaceae

[Sarcolaenaceae + Dipterocarpaceae]] [Cytinaceae + Muntingiaceae] Malvaceae]]

Brassicales

[Akaniaceae + Tropaeolaceae] [[Moringaceae + Caricaceae] [Setchellanthaceae

[Limnanthaceae [[Koeberliniaceae [Bataceae + Salvadoraceae]] [Emblingiaceae

[Pentadiplandraceae [Gyrostemonaceae Resedaceae Stixis and relatives] Tovariaceae

[Capparaceae [Cleomaceae + Brassicaceae]]]]]]]]

Berberidopsidales

Aextoxicaceae + Berberidopsidaceae

Santalales

Erythropalaceae Olacaceae [[Loranthaceae [Misodendraceae + Schoepfiaceae]]

[Opiliaceae + Santalaceae]] Balanophoraceae unplaced

Caryophyllales

[[Droseraceae [Nepenthaceae [Drosophyllaceae [Ancistrocladaceae +

Dioncophylleaceae]]]]] [[Frankeniaceae + Tamaricaceae] [Polygonaceae +

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

37

Plumbaginaceae]]] [Rhabdodendraceae [Simmondsiaceae [[Asteropeiaceae +

Physenaceae] [[Caryophyllaceae [Achatocarpaceae + Amaranthaceae]]

[Stegnospermataceae [Limeaceae [[Lophocarpaceae [Barbeuiaceae [Aizoaceae

[Sarcobataceae Phytolaccaceae Nyctaginaceae]]]] [Molluginaceae [Halophytaceae

Montiaceae Basellaceae Didiereaceae [Talinaceae [Portulacaceae [Anacampseros etc +

Cactaceae]]]]]]]]]]]]]

ASTERIDS

Cornales

[Cornaceae + Nyssaceae] [Hydrangeaceae + Loasaceae] Hydrostachyaceae [Curtisiaceae

+ Grubbiaceae]

Ericales

[Balsaminaceae [Marcgraviaceae + Tetrameristaceae]] [[Polemoniaceae +

Fouquieraceae] Lecythidaceae [[Sladeniaceae + Pentaphylacaceae] [Sapotaceae

[Ebenaceae [Maesaceae [Theophrastaceae [Myrsinaceae + Primulaceae]]]]]

[Mitrastemonaceae Theaceae [Symplocaceae [Styracaceae + Diapensiaceae]]

[[Sarraceniaceae [Roridulaceae + Actinidiaceae] [Clethraceae [Cyrillaceae +

Ericaceae]]]]]]

LAMIIDASTERID I

Unplaced

Metteniusaceae Oncothecaceae Icacinaceae and some unplaced ex-Icacinaceae

Garryales

Garryaceae + Eucommiaceae

Unplaced

Boraginaceae Vahliaceae

Gentianales

Rubiaceae [Gentianaceae [Loganiaceae [Gelsemiaceae + Apocynaceae]]]

Lamiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

38

Plocospermataceae [Oleaceae [Tetrachondraceae [Calceolariaceae

[Gesneriaceae Peltanthera [Pedaliaceae Carlemanniaceae Martyniaceae Stilbaceae

Plantaginaceae Linderniaceae Bignoniaceae Verbenaceae Lamiaceae Paulowniaceae

Schlegeliaceae Thomandersiaceae Phrymaceae Rehmannia Acanthaceae

Scrophulariaceae Orobanchaceae Byblidaceae Lentibulariaceae]]]]]

Solanales

[Montiniaceae [Sphenocleaceae + Hydroleaceae]] [Convolvulaceae + Solanaceae]

CAMPANULIDASTERID II

Aquifoliales

[[Cardiopteridaceae + Stemonuraceae] [Aquifoliaceae [Helwingiaceae +

Phyllonomaceae]]

Asterales

[Rousseaceae + Campanulaceae] Pentaphragmataceae [[Alseuosmiaceae [Phellinaceae

+ Argophyllaceae]] Stylidiaceae [Menyanthaceae [Goodeniaceae [Calyceraceae +

Asteraceae]]]]

Escalloniales

Escalloniaceae

Bruniales

[Bruniaceae + Columelliaceae]

Apiales

Pennantiaceae [Torricelliaceae [Griseliniaceae [Pittosporaceae [Araliaceae

[Myodocarpaceae + Apiaceae]]]]]

Paracryphiales

Paracryphiaceae

Dipsacales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

39

Adoxaceae [Diervillaceae [Caprifoliaceae [Linnaeaceae [Morinaceae [Dipsacaceae +

Valerianaceae]]]]]

LINKS TO ORDERS AND FAMILIES

[Back to Top]

MAIN GROUPINGS

Asterids commelinids core eudicots asterid 1 asterid

2 eudicots gymnosperms Magnoliophyta monocots N-fixing claderosids seed plants

ALPHABETICAL LISTING OF ALL ORDINAL NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | L | M | N | O | P | Q | R | S | T | U | V | W | X | Z

Abietales Acanthales Acerales Acorales Actinidiales Actinostrobales Adoxales Aescul

ales Agavales Aizoales AkanialesAlismatales Alliales Alseuosmiales Alstroemeriales

Altingiales Amaranthales Amaryllidales Amborellales AmbrosialesAmmiales Amomal

es Ancistrocladales Anisophylleales Annonales Anthobolales Apiales Apocynales Apo

nogetonalesAquifoliales Arales Araliales Aralidiales Araucariales Arecales Aristolochi

ales Athrotaxidales Asarales AsclepiadalesAsparagales Asphodelales Asparagales Ast

eliales Atriplicales Aucubales Austrobaileyales Avenales

Balanitales Balanopales Balanophorales Balsaminales Barbeyales Barclayales Batales

Begoniales BerberidalesBerberidopsidales Betulales Biebersteiniales Bignoniales Bix

ales Boraginales Brassicales Brexiales Bromeliales BrunialesBrunoniales Burmannial

es Burserales Butomales Buxales Byblidales

Cactales Callitrichales Calycanthales Calycerales Campanulales Campynematales Can

ellales Cannales CapparalesCaprifoliales Cardiopteridales Carduales Caricales Carle

manniales Caryophyllales Cassiales Casuarinales CelastralesCentrolepidales Cephalot

ales Cephalotaxales Ceratophyllales Cercidiphyllales Chenopodiales Chironiales Chlor

anthalesChrysobalanales Cinchonales Circaeasterales Cistales Citrales Cocosales Col

chicales Columelliales CombretalesCommelinales Connarales Convolvulales Coriarial

es Cornales Corylales Corynocarpales Crassulales CrossosomatalesCucurbitales Cunn

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

40

inghamiales Cunoniales Cupressales Cycadales Cyclanthales Cymodoceales Cynarales

CynomorialesCyperales Cyrillales Cytinales

Daphnales Daphniphyllales Dasypogonales Datiscales Degeneriales Desfontainiales D

iapensiales Didymelales DillenialesDioales Dioncophyllales Dioscoreales Diospyrales

Dipentodontales Dipsacales Droserales

Ebenales Echiales Elaeagnales Elaeocarpales Elatinales Elodeales Emmotales Empetr

ales Ephedrales Ericales EriocaulalesErythropalaless Escalloniales Eucommiales Euph

orbiales Eupomatiales Eupteleales Euryalaless

Fabales Fagales Falcatifoliales Ficales Flacourtiales Flagellariales Fouquieriales Franc

oales Frangulales

Galiales Garryales Geissolomatales Gentianales Geraniales Gesneriales Ginkgoales G

laucidiales Globulariales GnetalesGoodeniales Greyiales Griseliniales Grossulariales

Grubbiales Gunnerales Gyrocarpales Gyrostemonales

Haemodorales Haloragales Hamamelidales Hanguanales Heisteriales Helleborales He

lwingiales Himantandrales HippuridalesHomaliales Hortensiales Huales Huerteales

Hydatellales Hydnorales Hydrangeales Hydrastidales HydrocharitalesHydropeltidales

Hydrostachyales Hypericales Hypoxidales

Icacinales Illiciales Iridales Irvingales Iteales Ixiales

Jasminales Juglandales Julianiales Juncaginales Juncales

Lacistematales Lactoridales Lamiales Lardizabalales Laurales Lecythidales Ledocarpal

es Leitneriales LentibularialesLigustrales Liliales Limnanthales Linales Liriales Loasal

es Lobeliales Loganiales Lonicerales Loranthales LowialesLythrales

Magnoliales Malpighiales Malvales Marathrales Marcgraviales Mayacales Medusagy

nales Medusandrales MelanthialesMelastomatales Meliales Melianthales Meliosmal

es Menispermales Menyanthales Metteniusales MitrastemonalesMiyoshiales Moni

miales Moringales Musales Myricales Myristicales Myrothamnales Myrsinales Myrta

les

Najadales Nandinales Narcissales Nartheciales Nelumbonales Nepenthales Neuradal

es Nitrariales Nolanales NothofagalesNyctaginales Nymphaeales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

41

Ochnales Oenotherales Olacales Oleales Onagrales Oncothecales Opuntiales Orchid

ales Oxalidales

Saxifragales Pandales Pandanales Papaverales Paracryphiales Parasitaxales Paridales

Parnassiales PassifloralesPenaeales Pennantiales Pentaphragmatales Peridiscales P

etiveriales Petrosaviales Phellinales Philydrales PhyllanthalesPhysenales Phytolaccal

es Picramniales Pinales Pinguiculales Piperales Pittosporales Plantaginales Platanale

sPlumbaginales Poales Podocarpales Podophyllales Podostemales Polemoniales Pol

ygalales Polygonales PontederialesPortulacales Posidoniales Potamogetonales Primu

lales Proteales

Quercales Quillajales Quintiniales

Rafflesiales Ranunculales Rapateales Resedales Restionales Rhabdodendrales Rhamn

ales Rhinanthales RhizophoralesRhodorales Rhoipteleales Roridulales Rosales Rouss

eales Rubiales Ruppiales Rutales

Sabiales Salicales Salvadorales Samolales Samydales Sanguisorbales Santalales Sapin

dales Sapotales SarracenialesSaxegotheales Saxifragales Scheuchzeriales Sciadopitya

les Scleranthales Scrophulariales Scyphostegiales SedalesSilenales Simmondsiales S

milacales Solanales Sphenocleales Sphenostemonales Stangeriales Stellariales Stemo

nalesStilbales Stylidiales Styracales Surianales

Taccales Tamales Tamaricales Taxales Taxodiales Tecophilaeales Terebinthales Tern

stroemiales Theales TheligonalesThymelaeales Tiliales Tofieldiales Torricelliales Tov

ariales Tribelales Trilliales Trimeniales Triuridales TrochodendralesTropaeolales Turn

erales Typhales

Ulmales Urticales

Vacciniales Vahliales Vallisneriales Velloziales Veratrales Verbenales Viburnales Vinc

ales Violales Viscales VitalesVochysiales

Welwitschiales Winterales

Xanthorrhoeales Ximeniales Xyridales

Zamiales Zingiberales Zosterales Zygophyllales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

42

ALPHABETICAL LISTING OF ALL FAMILY NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | K | L | M | N | O | P | Q | R | S | T | U | V | W | X | Y

|Z

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In some cases clicking on a family synonym will send you directly to the subfamily in

which the genus of the basionym is to be found

Abietaceae Abolbodaceae Abrophyllaceae Acalyphaceae Acanthaceae [Acanthochla

mydaceae] Acarnaceaee AceraceaeAchariaceae Achatocarpaceae Achradaceae Achy

ranthaceae Acmopylaceae Acoraceae Acristaceae ActaeaceaeActinidiaceae Actinota

ceae Amentotaxaceae Adenogrammaceae Adoxaceae Aegialitidaceae Aegicerataceae

Aegilopaceae Aeginetiaceaee Aegiphilaceae Aesculaceae Aextoxicaceae Agapanthac

eae Agavaceae AgdestidaceaeAgialidaceae Agrimoniaceae Agrostidaceae Ailanthace

ae Aitoniaceae Aizoaceae Akaniaceae Alangiaceae AlchemillaceaeAldrovandaceae A

lismataceae Alliaceae Allioniaceae Allophylaceae Aloaceae Alopecuraceae Alpiniacea

e AlseuosmiaceaeAlsinaceae Alsodeiaceae Alstroemeriaceae Altingiaceae Alzateacea

e Amaranthaceae Amaryllidaceae Amborellaceae Ambrosiaceae Amentotaxaceae A

mmanniaceae Ammiaceae Amomaceae Ampelidopsaceae Amygdalaceae Amyridacea

eAnacardiaceae Anagallidaceae Anarthriaceae Anchusaceae Ancistrocladaceae Andr

omedaceae AndropogonaceaeAndrostachyaceae Androsynaceae Anemarrhenaceae

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ridaceae Apostasiaceae Aptandraceae AquifoliaceaeAquilariaceae Aquilegiaceae A

raceae Aragoaceae Araliaceae Aralidiaceaee Araucariaceae Arbutaceae Arceuthidace

aeArceuthobiaceae Arctostaphylaceae Arctotidaceae Ardisiaceae Arecaceae Argoph

yllaceae Arisaraceae AristolochiaceaeAristoteliaceae Arjonaceae Armeriaceae Artem

isiaceae Arthrotaxidaceae Artocarpaceae Arundinaceae ArundinellaceaeAsaraceae A

sclepiadaceae Ascyraceaee Asiraceae Asparagaceae Asperulaceae Asphodelaceae As

pidistraceaeeAsteliaceae Asteraceae Asteranthaceae Asteropeiaceae Astragalaceae

Astrocarpaceae AthanasiaceaeAtherospermataceae Atriplicaceae Atropaceae Aucuba

ceae Aurantiaceae Austrobaileyaceae Austrotaxaceae AvenaceaeAverrhoaceae Avet

raceae Avicenniaceae Azaleaceae Azimaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

43

Balanitaceae Balanopaceae Balanophoraceae Balsameaceae Balsaminaceae Bambusa

ceae Barbaceniaceae BarbeuiaceaeBarbeyaceae Barclayaceae Barringtoniaceae Base

llaceae Bataceae Baueraceae Bauhiniaceae Baxteriaceae BegoniaceaeBehniaceae B

elangeraceae Belloniaceae Bembiciaceae Berberidaceae Berberidopsidaceae Berryac

eaee BersamaceaeBertyaceae Berzeliaceae Besleriaceae Betaceae Betulaceae Biebe

rsteiniaceae Bifariaceae Bignoniaceae BischofiaceaeBixaceae Blakeaceae Blandfordia

ceae Blattiaceae Blepharocaryaceae Blitaceae Blyxaceae Boerlagellaceae Bolivaracea

eBombacaceae Bonnetiaceae Bontiaceae Boopidaceae Boraginaceae Borassaceae B

oroniaceae BoryaceaeBotryodendraceae Bougainvilleaceae Boweniaceae Brassicacea

e Brachycaulaceae Bretschneideraceae BrexiaceaeBromeliaceae Brownlowiaceae Br

unelliaceae Bruniaceae Brunoniaceae Brunsvigiaceae Bryoniaceae BuchneraceaeBuci

daceae Buddlejaceae Buglossaceae Bulbocodiaceae Bumeliaceae Bupleuraceae Burc

hardiaceae BurmanniaceaeBurseraceae Butneriaceae Butomaceae Buxaceae Byblida

ceae Byttneriaceae

Cabombaceae Cacaoaceae Cactaceae Caesalpiniaceae Caladiaceae Calamaceae Calce

olariaceae CalectasiaceaeCalendulaceae Callaceae Callicomaceae Calligonaceae Calli

traceae Callitrichaceae Calochortaceae CalophyllaceaeCalthaceae Calycanthaceae Ca

lyceraceae Cambogiaceae Camelliaceae Campanulaceae CampynemataceaeCanacom

yricaceae Canellaceae Cannabaceae Cannaceae Canopodaceae Canotiaceae Cansjera

ceae CapparaceaeCaprariaceae Caprifoliaceae Cardiopteridaceae Carduaceae Carica

ceae Carissaceae Carlemanniaceae CarpinaceaeCarpodetaceae Cartonemataceae Ca

ryocaraceae Caryophyllaceae Casearieaceae Cassiaceae CassipoureaceaeCassythacea

e Castaneaceae Castelaceae Casuarinaceae Catesbaeaceae Cathedraceae Cecropiace

ae CedraceaeCedrelaceae Celastraceae Celosiaceae Celtidaceae Centaureaceae Cen

trolepidaceae Centroplacaceae CepaceaeCephalanthaceae Cephalotaceae Cephalota

xaceae Cerastiaceae Ceratoniaceae Ceratophyllaceae CerberaceaeCercidiphyllaceae

Cercocarpaceae Cercodiaceae Cereaceae Cerinthaceae Ceroxylaceae Cestraceae Cev

alliaceaeChailletiaceae Chamaedoraceae Chamaemoraceae Chamelauciaceae Chaun

ochitonaceae Chelidoniaceae ChelonaceaeChenopodiaceae Chimonanthaceae Chingi

thamnaceae Chionographidaceae Chiranthodendraceae ChironiaceaeChlaenaceae Ch

loanthaceae Chloranthaceae Chloridaceae Chlorogalaceae Chrysobalanaceae Cicerac

eae CichoriaceaeCimicifugaceae Cinchoniaceae Circaeaceae Circaeasteraceae Cissac

eae Cistaceae Citraceae Clematidaceae CleomaceaeClethraceae Clusiaceae Cneorac

eae Cnestidaceae Cnicaceae Cobaeaceae Cochlospermaceae Cocosaceae Coffeaceae

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Colchicaceae Coleogynaceae Columelliaceae Combretaceae Commelinaceae Comocl

adiaceae Compositae CompsoaceaeConantheraceae Connaraceae Conostylidaceae C

onvallariaceae Convolvulaceae Cordiaceae CoreopsidaceaeCoriandraceae Coriariacea

e Coridaceae Corispermaceae Cornaceae Corokiaceae Coronillaceae Correaceae Cor

rigiolaceaeCorsiaceae Corydalaceae Corylaceae Corynocarpaceae Coryphaceae Cost

aceae Cotyledonaceae CoulaceaeCoutariaceae Coutoubiaceae Crassulaceae Crescen

tiaceae Cressaceae Crinaceae Crocaceae CroomiaceaeCrossosomataceae Crotonacea

e Cruciferae Cryptaceae Crypteroniaceae Cryptocorynaceae CryptomeriaceaeCtenol

ophonaceae Cucurbitaceae Cunninghamiaceae Cunoniaceae Cupressaceae Curcumac

eae Curtisiaceae CuscutaceaeCyananthaceae Cyanastraceae Cynanchaceae Cyanella

ceae Cycadaceae Cyclanthaceae CyclantheraceaeCyclocheilaceae Cydoniaceae Cymo

doceaceae Cynaraceae Cynocrambaceae Cynomoriaceae Cyperaceae CyphiaceaeCyp

hocarpaceae Cypripediaceae Cyrillaceae Cyrtandraceae Cyrtanthaceae Cytinaceae

Dacrycarpaceae Dactylanthaceae Damasoniaceae Daphnaceae Daphniphyllaceae Das

ypogonaceae DatiscaceaeDaturaceae Daucaceae Davidiaceae Davidsoniaceae Decais

neaceae Deeringiaceae Degeneriaceae DelphiniaceaeDendrophthoaceae Columelliac

eae Detariaceae Dialypetalanthaceae Dianellaceae Dianthaceae DiapensiaceaeDicha

petalaceae Dichondraceae Diclidantheraceae Dicrastylidiaceae Dictamnaceae Didiere

aceae DidymelaceaeDidymocarpaceae Diegodendraceae Diervillaceae Digitalidaceae

e Dilatridaceae Dilleniaceae Dioaceae DionaeaceaeDioncophyllaceae Dioscoreaceae

Diosmaceae Diospyraceae Dipentodontaceae Diphylleiaceae Diplolaenaceae Dipsaca

ceaeDipterocarpaceae Dirachmaceae Disanthaceae Diselmaceae Dodonaeaceae Do

mbeyaceae Donatiaceae DorsteniaceaeDortmannaceae Doryanthaceae Drabaceae D

racaenaceae Dracontiaceae Drimyidaceae Droseraceae DrosophyllaceaeDryadaceae

Duabangaceae Duckeodendraceae Dulongiaceae Durantaceae Durionaceae Dysphani

aceae

Ebenaceae Eccremidaceae Ecdeiocoleaceae Echiaceae Echinopaceae Ehretiaceae Ela

eagnaceae ElaeocarpaceaeElegiaceae Elatinaceae Ellisiaceae Ellisiophyllaceae Elodea

ceae Elytranthaceae Embeliaceae Emblingiaceae EmmotaceaeEmpetraceae Encepha

lartaceae Engelhardtiaceae Enhalaceae Epacridaceae Ephedraceae Ephemeranthacea

eEpilobiaceae Epimediaceae Eragrostidaceae Eremolepidaceae Eremosynaceae Erica

ceae Erinaceae EriocaulaceaeEriogonaceae Eriospermaceae Erodiaceae Erycibaceae

Eryngiaceae Erysimaceae Erythroniaceae ErythropalaceaeErythrospermaceae Erythro

xylaceae Escalloniaceae Eschscholziaceae Eucomidaceae Eucommiaceae Eucryphiace

aeEuonymaceae Eupatoriaceae Euphorbiaceae Euphrasiaceae Euphroniaceae Eupo

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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matiaceae Eupteleaceae EuryalaceaeEustrephaceae Euthemidaceae Eutocaceae Exb

ucklandiaceae Exocarpaceae

Fabaceae Fagaceae Falcatifoliaceae Ferulaceae Festucaceae Ficaceae Ficoidaceae Fit

zroyaceae FlacourtiaceaeFlagellariaceae Flindersiaceae Foetidiaceae Forestieraceae

Fothergillaceae Fouquieriaceae Fragariaceae FrancoaceaeFrangulaceae Frankeniacea

e Fraxinaceae Fraxinellaceae Freycinetiaceae Fritillariaceae Fuchsiaceae Fumariacea

eFunkiaceae

Gaiadendraceae Galacaceae Galanthaceae Galaxiaceae Galedupaceae Galeniaceae G

aliaceae Garciniaceae GardeniaceaeGardneriaceae Garryaceae Geissolomataceae Ge

itonoplesiaceae Gelsemiaceae Geniostomaceae GentianaceaeGeonomataceae Geosir

idaceae Geraniaceae Gerrardinaceae Gesneriaceae Gethyllidaceae Gilliesiaceae Gina

lloaceaeGinkgoaceae Gisekiaceae Gladiolaceae Glaucidiaceae Glechomaceae Glinace

ae Globulariaceae Gnaphaliaceae GnetaceaeGoetzeaceae Gomortegaceae Gomphiac

eae Gomphrenaceae Gonystylaceae Goodeniaceae Gordoniaceae GouaniaceaeGoup

iaceae Gramineae Gratiolaceae Grewiaceae Greyiaceae Grielaceae Griseliniaceae Gr

onoviaceae GrossulariaceaeGrubbiaceae Guaiacanaceae Guamatelaceae Guettardac

eae Gunneraceae Gustaviaceae Guttiferae GyrocarpaceaeGyrostemonaceae

Hachetteaceae Haemanthaceae Haemodoraceae Halesiaceae Hallieraceae Halocarpa

ceae Halophilaceae HalophytaceaeHaloragaceae Hamamelidaceae Hanguanaceae H

aptanthaceae Harmandaceae Hebenstretiaceae HectorellaceaeHederaceae Hedyosm

aceae Hedyotidaceae Hedysaraceae Heisteriaceae Heleniaceae Heliamphoraceae H

elianthaceaeHelianthemaceae Helichrysaceae Heliconiaceae Helicteraceae Heliotropi

aceae Helleboraceae Heloniadaceae HelosidaceaeHelwingiaceae Hemerocallidaceae

Hemimeridaceae Hemiphylacaceae Henriqueziaceae Henslowiaceae Hermanniaceae

Hernandiaceae Herniariaceae Herreriaceae Hesperocallidaceae Heterantheraceae He

teropyxidaceae HeterostylaceaeHibbertiaceae Hibiscaceae Hilleriaceae Himantandra

ceae Hippocastanaceae Hippocrateaceae HippomanaceaeHippophaeaceae Hippurida

ceae Hirtellaceae Holacanthaceae Homaliaceae Hopkinsiaceae Hoplestigmataceae H

ordeaceaeHornschuchiaceae Hortensiaceae Hortoniaceae Hostaceae Houstoniaceae

Huaceae Huerteaceae HugoniaceaeHumbertiaceae Humiriaceae Hyacinthaceae Hyd

atellaceae Hydnoraceae Hydrangeaceae Hydrastidaceae HydrillaceaeHydrocharitacea

e Hydrocotylaceae Hydrogetonaceae Hydroleaceae Hydropeltidaceae Hydrophylacac

eae HydrophyllaceaeHydrostachyaceae Hymenocardiaceae Hypecoaceae Hyperanth

eraceae Hypericaceae Hypopityaceae HypoxidaceaeHypseocharitaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta

ceae ImperatoriaceaeInocarpaceae Inulaceae Ionidiaceae Iriarteaceae Iridaceae Irvi

ngiaceae Isnardiaceae Isophysidaceae IteaceaeStrasburgeriaceae Ixiaceae Ixioliriace

ae Ixonanthaceae

Jalapaceae Jamboliferaceae Japonoliriaceae Jasionaceae Jasminiaceae Johnsoniaceae

Joinvilleaceae JuglandaceaeJulianiaceae Juncaceae Juncaginaceae Juniperaceae Jus

siaeaceae Justiciaceae

Kadsuraceae Kaliphoraceae Kalmiaceae Kaniaceae Kiggelariaceae Kingdoniaceae Kin

giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae

Krameriaceae

Labiatae Lacandoniaceae Lachenaliaceae Lacistemataceae Lactoridaceae Lactucaceae

Lagerstroemiaceaee LamiaceaeLampsanaceae Lanariaceae Langsdorffiaceae Lantan

aceae Lapageriaceae Lardizabalaceae Lasiopetalaceae LauraceaeLawsoniaceae Laxm

anniaceae Lecythidaceae Ledaceae Ledocarpaceae Leeaceae Leguminosae Leitneriac

eae LemnaceaeLennoaceae Lentibulariaceae Lentiscaceae Leoniaceae Leonticaceae

Lepidobotryaceae LepidocarpaceaeLepidocaryaceae Lepidocerataceae Leptaulaceae

Leptospermaceae Lepturaceae Lepuropetalaceae LeuctenbergiaceaeLibocedraceae

Licaniaceae Ligustraceae Lilaeaceae Liliaceae Limeaceae Limnanthaceae Limnocharit

aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace

ae Linderniaceae Lindleyaceae Linnaeaceae LippayaceaeLiriaceae Liriodendraceae Li

ssocarpaceae Littorellaceae Loasaceae Lobeliaceaee Loganiaceae LomandraceaeLoni

ceraceae Lopeziaceae Lophiocarpaceae Lophiolaceae Lophiraceae Lophophytaceae L

ophopyxidaceae LoranthaceaeLotaceae Lowiaceae Lupulaceaee Luxemburgiaceae Lu

zuriagaceae Lychnidaceae Lyciaceae LyginiaceaeLygodisodeaceae Lysimachiaceae Ly

thraceae

Macarisiaceae Mackinlayaceae Madiaceae Maesaceae Magnoliaceae Malaceae Male

sherbiaceae MalortieaceaeMalpighiaceae Malvaceae Mangiaceae Manicariaceae Ma

paniaceae Marantaceae Marathraceae MarcgraviaceaeMartyniaceae Mastixiaceae

Matricariaceae Maundiaceae Mayacaceae Medeolaceae Medusagynaceae Peridiscac

eaeMelaleucaceae Melampyraceae Melanophyllaceae Melanthiaceae Melastomatac

eae Meliaceae Melianthaceae MelicaceaeMeliosmaceae Melittidaceae Melochiacea

e Memecylaceae Mendonciaceae Menispermaceae Menthaceae MenyanthaceaeMe

nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

47

Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

Nypaceae Nyssaceae

Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

ceae Papilionaceae Pappophoraceae PapyraceaeParacryphiaceae Parasitaxaceae Par

ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

48

hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

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cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 2: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

2

INTRODUCTION

This linked series of four pages is a simplified version of the main site The focus is on

the larger patterns evident in the phylogeny evolution and diversification of seed

plants and on morphological variation in general

Systematics is a profoundly historical discipline and we forget this at our peril Only

with a phylogeny can we begin to understand diversification regularities in patterns of

evolution or simply suggest individual evolutionary changes within a clade Our

recovery of that phylogeny is the recovery of evidence of a series of unique events that

comprises the history of life Although our knowledge of the major clades of seed plants

and the relationships within and between them are still somewhat in a state of flux

much of the broad outline is clear (see the Angiosperm Phylogeny Group II 2003)

Furthermore as details of phylogeny are clarified and new findings made in anatomy

morphology etc they can be rapidly integrated the Angiosperm Phylogeny Group

system that is followed here Books are out-of-date before they appear furthermore

there is no comprehensive phylogeny-based treatment of angiosperms out-of-date or

not D Soltis et al (2005b) is the closest (also take the following link to a regularly

updated and printable Angiosperm Phylogeny Poster)

These pages are for undergraduate students with an interest in botany and some

knowledge of basic plant morphology and for those with a general interest in seed plant

evolution They are made up of a much modified Apomorphies page that has been

combined with the order pages all very much changed The focus is on providing

possible apomorphies for as many clades as possible although there is much less detail

than in the Apomorphies page (much chemistry details of ovule morphology other than

general appearance most indications of base chromosome number etc have been

removed) and I have simplified the vocabulary somewhat There is also information on

the sizes and distributions of families lists of the larger genera included features by

which families can be recognised comments about the ages (to be treated with

considerable caution) and diversification of groups and about the major herbivores

pollinators and seed dispersers associated with them notes on interesting

morphological variation summaries of what is known about infrafamilial phylogenetic

relationships etc The literature cited will allow the student find the major

morphological taxonomic and phylogenetic information about each family especially

when the Characters page is also consulted Phylogenies of most orders and of some

families (eg Poaceae) are included but not maps or images for these there are links

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

3

from each family to the more detailed treatments on the individual order pages In

larger families I tend to focus on literature that deals with monophyletic groups that

include fifty or more taxa in smaller families the coverage is more detailed

I have emphasized plant families because they are the groups - admittedly partly

arbitrary as to circumscription but now for the most part monophyletic - around which

many of us organize our understanding of plant diversity I also pay attention to

groupings of families because so much progress has been made in the last decade in

particular in sorting them out Infrafamilial groups in groups like Poaceae Malvaceae

and Ericaceae are also included and these are being added to as studies become

available

But we dont want to know just about clades we want to know what makes clades

unique the synapomorphies or shared derived characters of those clades that first

appeared in their immediate ancestors However for the most part our knowledge of

synapomorphies remains poor ss we will see finding out the composition of clades is

often easier than finding the synapomorphies for the same clades (see the discussion

below) And of course knowing about synapomorphies is just one aspect of

understanding the whys and wherefores of the evolution and diversification of seed

plants our ultimate goal

ON CLASSIFICATIONS IN GENERAL AND THIS CLASSIFICATION IN PARTICULAR

On classifications in general

Classifications in the broad sense are box-in-box group-in-group or partwhole

naming devices that we use to communicate aspects of our knowledge of things in

general From this point of view there is nothing particularly special about biological

classifications apart perhaps from their sizes For any biological classification system to

be effective it must be stable universal (ie be used by a wide range of people)

comprehensive (if too many taxa are unplaced at the level of interest the classification

is of less utility) and it must enhance communication of knowledge by helping us to

relate things (Stevens 2006a for references) Phylogenetic classifications convey aspects

of our knowledge about phylogenetic relationships of organisms Thus a family is clearly

flagged as such and is a monophyletic group that can contain several genera also

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

4

flagged as such and also monophyletic but a genus can never include families Generic

family etc names are simply words we use to denote appropriate parts of phylogenies

and minimal aspects of their relationships and the irreducible aspect of relationships

that is emphasized here is monophyly A monophyletic group is one containing only and

all descendents of a common ancestor and it is often characterizable by apomorphies

that is shared derived characters that appeared in the ancestral lineage or stem clade

of that monophyletic group

Thus I am using a flagged ie ranked hierarchy for naming taxa rather than the

unranked systems that have recently been proposed (see below) The rank terminations

used (-ales -aceae etc - the flags) merely suggest relative positions of groups in the

local hierarchy If Ericaceae and Vaccinioideae are part of the same monophyletic group

the latter must refer to a clade nested within the former even if neither can necessarily

be directly compared with Polemoniaceae and Cobaeoideae (other than all being

putatively monophyletic groups) Such a flagged hierarchy is useful as a mnemonic and

communication device (eg Stevens 2006a) It improves memorization and emphasis on

families and orders as here is a didactic device - families are monophyletic units useful

in communication major units learned by biologists and others world-wide

The distinction between grouping and ranking is extremely important as is how we

interpret the latter We can both agree that there is a genus Acer yet disagree as to

whether it should be in Aceraceae or submerged in Sapindaceae Although from one

point of view this disagreement is utterly trivial it can have profound consequences if

we misunderstand the nature of the classificatory hierarchy Taxa at the same rank are

equivalent only by designation and have nothing necessarily in common (unless they

are sister taxa) other than their monophyly Rank as used here has no meaning other

than signifying a monophyletic group that includes other monophyletic groups with

appropriately subordinate rank terminations Taxa at the same rank have often been

treated incorrectly as if they were equivalent by biologists attempting to understand

evolutionary or biogeographic problems (see Bertrand et al 2006 for detailed

discussion) even if those constructing or using classifications - including Darwin (1859) -

have been explicit about the non-equivalence of taxa at the one rank (Stevens 1997) In

fact rank terminations have relatively infrequently been used by taxonomists to reflect

absolute rank although Linnaeus (at least in theory) at the level of genus and species

may be such an example (Classifications where rank is absolute taxa at the same rank

somehow being comparable entities are class hierarchies in the strict sense - Stevens

2002 2006a) It has also been suggested that taxon rank be adjusted so that rank

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

5

somehow reflects the degree of morphological differences between taxa or that taxa at

the same rank be based on similar characters or show a similar amount of distinctness

This might be possible using phenetic methods of analysis but is very difficult if ones

classification is phylogeny-based as here it could promote instability if used in taxa

where such a criterion had not previously been used and it might also inadvertently

suggest that taxa might be equivalent (for an example see Fritsch et al 2008) There

have also been proposals that rank could reflect the age of the clade (eg Hennig 1966)

with clades that have diverged by a particular time all being given the same rank Apart

from the fact that aging times of divergence of clades is still a difficult enterprise huge

disruptions to our nomenclature would result Recent suggestions which invoke the use

of age in classifications focus on providing a standardized timeclip ie a set of letters

referring to a particular geological period that could simply be added to a conventional

taxon name (Avise amp Mitchell 2007) However even such timeclips are unlikely to come

into general use soon

It would be impossible even to think about a higher-level classification such as this

without the advances in our understanding of relationships made by the phylogenetic

analyses of molecular data carried out over the last twenty years One can then

integrate the data to be found in both classical and recent morphological studies with

these phylogenies For the dramatic changes in this area see for instance the

pessimistic attitude towards orders in Davis and Heywood (1963 107-108) The most

unsatisfactory taxon in Angiosperm classification they were indefinable their

circumscription was not fixed etc Families they thought were likely to be the largest

natural unit within the mono- or dicotyledons Along the same lines almost three

quarters of the orders (4459 monofamilial orders ignored) recognised by Cronquist

(1981) are not monophyletic ie they do not contain all and only the descendents of a

common ancestor Most of those orders that are monophyletic are very small

(Zingiberales with eight families are the largest) for families on the other hand about

two thirds (189273) are monophyletic

Turning now to phylogenetic classifications and the particular classification used

here Backlund and Bremer (1998) provide a useful discussion on the principles of

phylogenetic classification that is applicable at all levels apart from species (see also

Stevens 1998 also Albach et al 2004 Entwisle amp Weston 2005 Pfeil amp Crisp 2005 etc

for examples) Backlund and Bremers main principle is that taxa that are recognised

formally should be monophyletic However this does not indicate which particular

clades we might wish to name as families genera etc and talk about in general

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

6

conversation If a well-supported hypothesis of monophyly is a necessary prerequisite

for a group to be named it is not a sufficient prerequisite (but cf the PhyloCode -

Cantino amp de Queiroz 2006 Cantino et al 2007) Not all clades need be named indeed

it would barely be practicable (or practical) to do this To decide which clades should be

named additional criteria can be invoked Other things being equal it is helpful if 1 taxa

formally recognised are easily recognizable 2 groups that are well-established in the

literature are preserved 3 the size of groups is taken into account and 4

nomenclatural changes are minimized (Backlund amp Bremer 1998) Thus numerous small

groups have little to recommend them since individually they summarize little

information and tend to clog the memory while groups that are too big may be

amorphous Somewhat similarly Godfray and Knapp (2004 p 562) note that users

want stable informative and accessible classifications that enable easy identification

(see also Simon 2008) - although invoking users without specifying those who make up

this group is not very helpful This classification is for all interested in comparative

biology hence the emphasis on monophyly Although the clades named are sometimes

difficult to characterise there are as we shall see many ways of making such a

classification accessible to all

Problems with this emphasis on monophyly may be caused by reticulation events

such as hybridization endosymbiosis and lateral gene transfer but they are unlikely to

be common confusing factors here Genera can often be pegged to above the level at

which hybridization is at all common However in Poaceae-Pooideae-Triticeae there are

some intractable problems where extremely well established common usage and the

principle of monophyly are likely to remain at odds Many genera are certainly not

monophyletic here being allopolyploids and the genera are ultimately based on the

different genomes they contain (Dewey 1984 Loumlve 1984 Barkworth 2000 for a history

of Triticeae classification Petersen et al 2006) There is also extensive reticulation

reported within Danthonioideae (Pirie et al 2009) Evidence also increases of old

hybridization events elsewhere in flowering plants that at the very least cause

discordance between relationships suggested by different genomic compartments as in

Smedmark and Anderberg (2007 Sapotaceae) and Fehrer et al (2007) Morgan et al

(2009) and Pelser et al (2008 all Asteraceae - hybridization is likely to be quite a

problem at the generic level here) and genera like Medicagoare turning out to have

highly reticulating relationships at the species level (Maureira-Butler et al 2008) Of

course there are many problems trying to apply the concept of monophyly to species

and for many - but not all - biologists strict monophyly is a less important criterion at

this level (Funk amp Omland 2003 see much of the discussion in Houmlrandl 2006)

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

7

The major endosymbiotic events that characterize the clade of which flowering

plants are a part (and gave rise to chloroplasts and mitochondria) are very ancient and

cause no problems for the student of multicellular organisms However lateral gene

transfer has been detected in a number of situations between quite unrelated

organisms (eg Bergthorsson et al 2003 - Amborella and liverworts see also below but

cf Goremykin et al 2009) and it may be particularly common in mitochondria (Sanchez-

Puerta et al 2008) Here too there are no major problems providing one is careful

such transfers do however raise all sorts of interesting biological questions (see

Richardson amp Palmer 2007 for a summary) Although there is increasing evidence for the

importance of genome duplications - hybridization is one cause of this - at various times

during the evolution of seed plants and of palaeopolyploidy events within eg the

Lauraceae and Magnoliaceae clades (Soltis et al 2009 for a summary see eg

the Characters page for further discussion) these too do not currently seem to pose

problems for the adoption of monophyly as the sine qua non of groups to be recognised

formally in this phylogenetic classification but it can make detecting orthologous genes

difficult

The accessory principles of Backlund and Bremer (1998) should be used in

combination Thus keeping the monogenericPlatanaceae separate from its sister

taxon Proteaceae is justifiable Both are much-used names that signal well supported

well defined and easily recognisable groups that have long been recognised as distinct

have several synapomorphies and do indeed look very unlike each other Combining

the two would yield a clade with few obvious apomorphies not to mention the fact

that Nelumbonaceae should by the same logic (it is also monogeneric) also be included

in the expanded family On the other hand it is difficult to justify the continued

recognition of Callitrichaceae or Hippuridaceae monophyletic and distinctive although

they may be If they were recognised several poorly characterised clades would also

have to be carved out ofPlantaginaceae in any classification that aimed to convey a

comprehensive view of the worlds flora The continued recognition

of Valerianaceae and Dipsacaceae also tends to run into this problem (see also Pfeil amp

Crisp 2005 Orthia et al 2005 Albach 2008 etc for useful practical discussions of such

matters) But there are no absolute guidelines If Podostemaceae turn out to be sister

to Hypericaceae (for references here and elsewhere in the Introduction see the

individual families) the subsequent moderate dismemberment of Clusiaceae sl is not

be too high a price to pay for the continued recognition of Podostemaceae Hence the

somewhat provisional recognition of Hypericaceae and Calophyllaceae as well as

Clusiaceae below the families can all be recognized and the name Podostemaceae in

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

8

particular is very well established In a somewhat similar situation if Lemna and its

relatives are a clade sister to most other Araceae should they be recognised as a

separate family Gymnostachys a phenetically fairly distinctive taxon as well as the less

phenetically distinct Orontioideae would have to be recognised as a separate families

(or combined as a single family) too but Araceae in a somewhat restricted sense would

be somewhat more morphologically coherent although not greatly so and not notably

distinct However it is in the very nature of such decisions to be somewhat arbitrary

and unsatisfactory hence the emphasis on consensus classifications here and on

classifications as simply being a means to an end

A useful distinction can be drawn between crown groups and stem groups The

former are monophyletic and include the extant members of a clade and their

immediate common ancestor (see the figure below) The groups characterized in this

site are such groups Thus Proteaceae here are crown group Proteaceae apomorphies

like the single carpel four-merous perianth etc being found in this common ancestor

Stem groups on the other hand include all the members of a lineage immediately after

its split from its sister group and all branches of this lineage In the case of Proteaceae

the stem group would include everything after its split from its sister group

Platanaceae Obviously most of the organisms in the lineage that terminates in crown-

group Proteaceae are unknown only a few being known as fossils and it is not known

exactly where particular apomorphies of crown group Proteaceae evolved along this

lineage

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

9

Indeed there is no reason other than convention or convenience why any group

should not be segregated into several smaller monophyletic groups or merged to

produce a larger unit we can talk about one large thing or about several smaller things

Thinking about aspects of size findings in ethnobiology and cognitive psychology can be

used to suggest that a moderate number - probably fewer than 500 - of families is a

reasonable goal at which to aim and that groupings of taxa throughout any system

should be rather small in size (eg Berlin 1992 Stevens 1994 1997) Major systems such

as those of Linnaeus and Bentham and Hooker were constructed explicitly so as to ease

the burden on the memory (Stevens 1997 2002 see also Scharf 2007) the latter in

particular ensuring that all groups in their classification were relatively small often

containing three to eight immediately subordinate taxa - but by no means all their

groups were formally named Along the same lines Burtt (1977b) suggested that the

number of names at any rank should be at most one third those at the immediately

lower rank - and monotypic taxa might not need a formal name Consistent with such

ideas a fairly broad view of families and orders is taken here whenever the constraints

of monophyly and other criteria used when constructing classifications (see above)

permit The APG system is thus a convention consistent with ideas such as those

expressed by Backlund and Bremer (1998) for similar conventions see eg the Grass

Phylogeny Working Group (2001) and for fungi Hibbett et al (2007)

There are other views Thus Takhtajan (1997) has suggested that smaller families

are more natural This is incorrect Monophyletic groups that include fewer taxa -

Takhtajans smaller families - do not necessarily have more apomorphies than larger

groups even if members of smaller groups will always have more features in general in

common than the larger group that includes them That is they will have their

apomorphies their unique features as well as progressively more plesiomorphies

features found both in the small groups and in the larger clades of which they are a part

having more features in common is indeed one common meaning of more natural

(Note that the implication of the word natural has long been a group of the kind

[usually unspecified] that I think should be recognised and if other botanists had

different ideas too bad their groups were necessarily not natural [see Bather 1927] It

is thus rarely a helpful word and is not used here) Furthermore if this approach is

adopted we will find a slippery slope ahead By this kind of argument all families should

be very small since their members will have a great deal in common and so will be

most natural However as families (for example) are split the relationships that are

evident between the segregates and that were responsible for their being placed in a

single family in the first place will seem to necessitate the recognition of a new order

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

10

etc as is evident in Takhtajans own work - general taxonomic inflation is the result (see

also comparable suggestions in a cladistic context for Brassicales in particular - Ronse de

Craene amp Haston 2006) Such splitting is also questionable when teaching and learning

families since the student needs to understand the system as a whole However for

some genera removed from the families that until now have included them the

phenetic-classificatory-phylogenetic structure in their new home may mandate the

recognition of small families On the other hand Takhtajans suggestion that narrowly

defined families are more useful for phylogenetic studies may be true Indeed I have

more than once regretted prematurely combining groups whether species (in the

context of monographic work) or families (in the course of preparing these notes) but

this is largely a separate issue

Van Steenis (1978) Philipson (1987b) and others have rightly questioned the value

of splitting a group when ideas of the relationships of its constituent members have not

changed - that is very good reasons have to be provided for splitting a family if the

genera within it remain part of the same clade rather than belonging to another clade

Thus APG (2003) broadened the circumscription of Malvaceae because of the

parapolyphyly of some of the families that had historically been associated with it (Judd

amp Manchester 1997 Alverson et al 1999 Bayer et al 1999) These families particularly

Tiliaceae and Sterculiaceae were not at all easy to distinguish their close relationship

(see eg Brown 1814) and overall similarity had long been conceded and to some

workers at least their combination has come as something of a relief Although most of

the larger clades within Malvaceae sl remain difficult to distinguish even with flowers

Cheek (2007) opts for a wholesale and novel dismemberment into ten families

however the very good reasons for doing this are wanting

The same principles are of course applicable when it comes to dividing genera little

other than a headache is gained by splitting genera such

as Drosera and Gnetum (Doweld 2000) as has recently been proposed Thus if an

established genus divides into two (or more) clades this is not a signal for recognising

two groups at the same level - so here it could be argued that the dismemberment

of Pterostylis (Jones amp Clements 2002b) was somewhat unfortunate Along the same

lines if a newly-discovered taxon is sister to an existing named taxon - say a genus - this

does not necessitate the description of a separate genus for the newly described species

(cf Davis 2002) (Of course some [eg Thorne 1976] have suggested that the sizes of

gaps between groups at the same rank should be similar but any principle like this is

inherently flawed since morphological gaps are more unstable than phylogenetic

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

11

relationships and applying it across all flowering plants would both be difficult and cause

substantial changes - see also below) Humphreys and Linder (2009) provide a well-

documented survey of generic concepts in plants which the reader should consult they

note that generic limits (broad versus narrow) have oscillated historically and that

currently larger genera tend to be recognised because studies tend to be on a broader

scale than in the past

Note that invoking similarity or difference - whether qualified (considerable

similarities substantial differences) or not - in a cladistic context as justification for

combining or splitting taxa is not a particularly strong argument (see eg

CardiopteridaceaeStemonuraceae - Karingrehed 2002c) Similarity and difference can

neither be defined precisely since what may seem to be substantial similarities to me

may not to the next person nor are they likely to be stable in the face of our changing

knowledge of morphology and what might be synapomorphies

I might have prefered to merge some families recognised here or split others but

by and large I do not think my own preferences matter very much - and I take the same

position with regards to comparable preferences expressed by others Indeed the

bottom line is that in flagged hierarchies of the kind used here the limits of any

monophyletic unit generally taught and discussed particularly other than species can

be established only by convention and consensus (eg Stevens 2002 2006a Entwisle amp

Weston 2005) This is rather different from the reasonable observation that the

phylogeny itself cannot be achieved by consensus (Thorne 1976) for in most cases there

is a fact of the matter when it comes to relationships Given the increasing support for

the outlines of angiosperm phylogeny a stable consensus classification based on this

phylogeny seems attainable Indeed in addition to providing current ideas of

relationships of seed plants in a synthesised form this site is part of an attempt to build

such a consensus about the circumscription of taxa (see APG 1999 2003 2009 Grass

Phylogeny Working Group 2001 Mabberley 2008) Reaching such a consensus is vital

since what we know of angiosperm phylogeny allows a very large number of

classifications to be based on it and as we find out more the number of possible

classifications increases greatly Unfortunately however nature does not dictate what

the classification should be All classifications are constructed by humans to

communicate particular aspects of groups and relationships Our goals as systematists

are surely to produce robust hypotheses of relationships to understand the evolution of

morphology and the like - but not to argue ad nauseam whether something should be

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

12

a family or a subfamily That way surely lies madness and worse the discredit of our

discipline

There are similar issues whatever naming system is used Thus in phylogenetic

naming (Baum et al 1998 for an example but cf Baum et al 2004 for the PhyloCode

see Cantino amp De Queiroz 2006) an unflagged hierarchy is used in which any

terminations of names used are uninformative about the relative position of taxa If one

adopts the principle of phylogenetic naming one indeed does not have to worry about

the nomenclatural consequences caused by lumping or splitting any well-supported

clade can be named without affecting the name of more or less inclusive clades

Unfortunately unflagged hierarchies have very serious deficiences as communication

devices because they lack one aspect essential in language biological or otherwise -

they contain no intrinsic information about the relationships of the group in question to

others (eg Pfeil amp Crisp 2005 Stevens 2006a) Recent suggestions for using prefixes like

Apo- and Pan- to PhyloCode names will however allow limited information of this

kind to be conveyed but only as it pertains to individual branches and current

proposals do not even mandate that the prefixes be employed consistently In any

event such proposals simply prevent the potential tripling of the number of quite

different names used to describe different aspects of a phylogenetic tree over those

used to name monophyletic groups pure and simple In general where n is the number

of extant species in a group the number of clades in such a group = n-1 (Species will

also need names too for their names see Dayrat et al 2008) Importantly here too

consensus over the clade names commonly learned by students and used in herbaria is

needed otherwise communication will be impeded the names themselves will provide

no guidelines as to which should be chosen The situation is of course more complicated

than this Terminations that convey ideas of rank in a phylogenetic classification can also

be used in phylocode names - however there they will carry no implications of rank

How they will be used is another matter of course

Of course there are other philosophies of classification and some still prefer

evolutionary classifications There classificatory principles differ substantially from those

followed here eg the recognition of paraphyletic taxa may be permitted however

detailed reasons for prefering the taxa that are recognised are rarely given although

nature and natural groups are often mentioned (cf Stuessy amp Koumlnig 2008) For

summaries of commonly used systems see Brummitt (1992) and Mabberley (2008)

new evolutionary systems appear every year or so However even those who allow or

promote the recognition of paraphyletic groups (eg Grant 2003 Thorne 2007

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

13

Heywood et al 2007) may find it of interest to examine a system recognizing only

monophyletic groups it provides a rather different understanding of evolution

Evolutionary classifications in general try and combine phylogeny and

morphological gaps although that is no easy thing to do - it is akin to combining chalk

and cheese (for an attempt to make this impossible task seem to be more objective see

Stuessy amp Koumlnig 2008) However there is a principle from evolutionary classification

that is relevant and I think quite useful here The size of the gap between two groups

tends to be inversely proportional to the sizes of the groups involved (Davis amp Heywood

1963) One can imagine a situation in which a large group is formally divided even

although the distinguishing characters of the two are weak whereas a smaller group

similarly divisable will be left intact

To summarize If hypotheses of phylogeny remain stable we should be able to base

a stable classification on that phylogeny and then get on with our work that is testing

the phylogenies we have elucidating phylogenies in areas where relationships are

unclear studying the evolution of morphology describing species etc In this context

the spread of the Angiosperm Phylogeny Group system (see below) and its widespread

utilisation in technical literature also floras (eg van der Meijden 2005) textbooks (eg

Simpson 2006 Judd et al 2007 [third edition]) dictionaries (Mabberley 2008) more

popular literature (eg Souza amp Lorenzi 2005 Spears 2006) and as an outline for a new

herbarium sequence (Haston et al 2007) is gratifying The posibility that one might be

able to develop a stable phylogeny-based classification of families and in particular

orders represents a dramatic turn-around from the pessimistic attitude about such

higher-level groupings expressed by Davis and Heywood (1963) and Thorne (1976) the

latter even suggesting that we should bury forever the metaphor of the phylogenetic

tree as highly unrealistic (ibid p 56) Returning to Godfray and Knapps (2004) users

of classifications who want a stable informative and accessible classification that

enables easy identification - unfortunately they want cake with everything and cannot

get it - these pages attempt to satisfy as many of their needs as possible but phylogeny

and monophyly are the primary shapers of the APG classification

On this classification in particular

Here I very largely follow the Angiosperm Phylogeny Group classification (APG

2003) Any differences are not to be interpreted as differences in principle simply that

new phylogenies continue to be published and here I attempt to provide an overview

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

14

of current ideas of higher-level relationships of seed plants The Angiosperm Phylogeny

Group classification is based on well-supported relationships evident in the numerous

molecular studies that began to appear in the late 1980s (see APG 1999 for the

principles underlying the classification) but recent phylogenetic work does not

contradict the major outlines of the trees used by APG II (2003) or even those in APG

I (APG 1999) The APG classification has for the most part been conservative and I

am similarly conservative here For the names and authorities I follow APG III (2009)

although the names of the authors may not always be correct these being in something

of a state of flux for the superordinal names used see Chase and Reveal (2009)

although classification at this level here is not exhaustive - only clades with several

orders have superordinal names

Some changes in our ideas of relationships and hence in the clades we talk about

are particularly likely in parts of Lamiales Caryophyllales and Malpighiales for example

although this will not affect the orders themselves Thus some changes to clade

circumscriptions are to be expected although I do not expect them to be substantial

but changes are neither a defect of cladistics nor a necessary consequence of the use of

molecular data Clades are hypotheses of relationships and as hypotheses they may be

overturned However work since APG I has not suggested other than minor changes

in the compositions of the orders even if the odd genus or even family is turning out to

be seriously misplaced - recent examples are Hydatellaceae (from monocots-Poales to

Nymphaeales Saarela et al 2007) Guamatelaceae (from Rosales-Rosaceae to

Crossosomatales Oh amp Potter 2006) and Perrottetia and Bhesa (from Celastrales-

Celastraceae to Huerteales and Malpighiales respectively Zhang amp Simmons 2006) The

main changes have been clarification of the relationhips of individual families or groups

of families that were of uncertain position eg of Chloranthaceae (Moore et al 2007)

Ceratophyllaceae (Jansen et al 2007) and unplaced asterid II families (Winkworth et al

2008a) As mentioned above phylogenies in the area of main interest in these pages are

overwhelmingly tree-like Although there is evidence for acquisition of host plant genes

by parasites (see below) even wider but inexplicable transfer of mitochochondrial

genes in plants like Amborella (Bergthorsson et al 2004 cf Goremykin et al 2009a)

and even transfers of nuclear genes (Vallenback et al 2008) these are the exception

rather than the rule

In cases where the Angiosperm Phylogeny Group II allowed alternatives as to the

limits of families - Papaveraceae in the broad sense or Papaveraceae plus

Pteridophyllaceae plus Fumariaceae Proteaceae in the broad sense or Proteaceae plus

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

15

Platanaceae - the choices made here follow common usage eg as in textbooks like

Judd et al (2007) and Simpson (2006) and particularly in the new edition of

Mabberleys The Plant Book (Mabberley 2008) This latter is itself an attempt to reflect a

consensus the result of taking the opinions of botanists at several meetings A largely

similar consensus is reflected in the APG III classification (APG 2009) where you will

see that alternative classifications have been dispensed with For many people the

existence of such alternative classifications will simply confuse so agreement over

which groupings to use when alternatives are permitted and then dispensing with the

alternatives that are not commonly used seems reasonable I am making the few

changes that are needed to bring the classification here in line with these others but

only gradually since there is not enough time in each day

As already mentioned higher-level relationships in general and the composition of

orders in particular have always presented something of a challenge to systematists

(eg Davis amp Heywood 1963) That being said the composition of some clades like

Apiales Crossosomatales and Pandanales is decidedly unexpected However it is

interesting to see that these higher level clades are generally accepted even in works

with different classificatory philosophies (eg Heywood et al 2007) For clades like

Malpighiales many of the family groupings within Asparagales etc attempts to find

distinctive characters have largely failed (but see Endress amp Matthews 2006a also

the Apomorphies page here) Interestingly as with families some groupings suggested

by molecular studies are supported by morphological andor chemical characters that

have long been known sometimes for over a hundred years the relationship between

Pittosporaceae and ApiaceaeAraliaceae are a case in point (Hegnauer 1969b and

references) As our knowledge of morphology and chemistry improves we can hope for

improvements in the characterisations of clades at all levels

In many cases the new family limits of the Angiosperm Phylogeny Group (see

APG 1999 2003) are not really controversial although changes from the limits

commonly accepted only a decade ago are sometimes quite dramatic (eg Wagenitz

1997) Thus the split of the old Saxifragaceae sl is necessitated by its extreme

polyphyly as also with Icacinaceaesl and Cornaceae sl However the limits and

relationships of such groups had long been considered to be unsatisfactory but until

recently there had been no real reason to prefer one proposed arrangement over

another It is generally accepted that the limits of Lamiaceae and Verbenaceae have to

be redrawn although there is as yet no compelling evidence that the redrawn taxa are

not sister taxa (there is no evidence that they are) But whatever their relationships the

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

16

content of the clades has changed considerably and incidentally they are now easier to

identify than before the decision to recognise two families is not difficult The same is

true for Salicaceae and Achariaceae (Malpighiales) two previously small families that

have received the bulk of the old Flacourtiaceae Clade and hence taxon limits remain

difficult around Theaceae (Ericalessee alsoSladeniaceae Pentaphylacaceae)

and Euphorbiaceae (Malpighiales see

also Phyllanthaceae Picrodendraceae Putranjivaceae etc) Here current groupings

may not represent quite such dramatic changes in our understanding of relationships

Indeed parts of the old Euphorbiaceae that were separated may yet go back together

particularly Phyllanthaceae and Picrodendraceae (see Wurdack et al 2004 Davis et al

2005) even if Theaceae should indeed be dismembered (Geuten et al 2004) However

given that there is no molecular evidence that currently warrants combining all the

segregates of Euphorbiaceae and even if some do come together the clades the

families represent suggest novel groupings not recognised in current classifications

Furthermore Rafflesiaceae appear to be embedded within Euphorbiaceae s str (Davis

et al 2007) so the family is divided Of course maintaining even a moderately narrowly-

circumscribed Euphorbiaceae would entail reducing the iconic Rafflesiaceae to

synonymy Relationships in core Caryophyllales especially

around Phytolaccaceae andNyctaginaceae and also Portulacaceae are incompletely

understood and refashioning of taxon limits will doubtless be needed as cladistic

relationships become apparent (eg see Nyffeler 2007 Ogburn 2007a b) Some

groupings in the old Icacinaceaeand Olacaceae also remain unclear

The discovery of the relationships of parasitic and aquatic groups have presented a

particular challenge to systematists Morphologically some of these plants are so highly

modified that interpretation of the plant body in conventional terms is difficult or even

impossible Thus parasitic groups (broad-leaved angiosperms only) such as Rafflesiaceae

are hard to place since both the vegetative body and the flowers are changed almost

beyond recognition (flowers of rafflesiaceae are a good example of gigantism - Davis et

al 2007) Furthermore plastid gene sequences may be difficult or impossible to obtain

the chloroplast DNA in particular being highly degraded and the rate of molecular

change in general may be high so the problem of long-branch attraction is serious (eg

Duff amp Nickrent 1997 Nickrent et al 1998 Caddick et al 2002a G Petersen et al

2006b) Echlorophyllous sapromycoheterotrophic taxa mostly known from monocots

present similar problems However progress is being made placements for

Apodanthaceae Rafflesiaceae Mitrastemonaceae Cytinaceae and Cynomoriaceae have

recently been suggested (Barkman et al 2004 2007 Davis amp Wurdack 2004 Nickrent et

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

17

al 2004 Davis et al 2007) it seems likely that Burmanniaceae sl are polyphyletic but

both parts are in Dioscoreales (Merckx et al 2006) and relationships within the largely

hemiparasitic Santalales are also gradually being clarified (Maleacutecot 2002) The inclusion

of parasitic taxa in general molecular analyses can cause conniptions (eg Nickrent et al

2004 Davis et al 2004 Chase et al 2006 G Petersen et al 2006b) and there can be

horizontal transmission of genes (eg Davis amp Wurdack 2005 Vitaceae to Rafflesiaceae

Barkman et al 2007 the mitochondrial atp1 gene commonly moves)

Similarly in water plants neither vessels in particular nor much xylem in general is

needed leaves are highly modified and water-mediated pollination if adopted may

well be associated with major changes in floral morphology Here too recent molecular

studies suggest that aquatic groups with hitherto problematic relationships may find

homes Thus Podostemaceae are close to Clusiaceae and Calophyllaceae and are sister

to Hypericaceae (Malpighiales Kita amp Kato 2001) Hydatellaceae which used to be in

Poales are part of Nymphaeales (Saarela et al 2007) and Hydrostachyaceae may be

close to Hydrangeaceae (Cornales Xiang et al 2002) or perhaps should be placed in

Lamiales note that in the first two cases in particular there are morphological and

chemical features that support such a move If Podostemaceae are indeed close to

Clusiaceae I look forward to seeing hypotheses to explain how the dramatic changes in

the vegetative body that have made Podostemaceae so problematic for generations of

systematists took place That conventional wisdom has trouble in understanding or

explaining how the morphologies of groups like Clusiaceae and Podostemaceae can be

related is largely a problem with conventional wisdom Ceratophyllaceae seem to be

finding a position as sister to eudicots (see Moore et al 2007) but their morphology is

so derived that there is no morphological evidence of which I am aware for this

relationship There are also similar although less extreme morphological problems with

many plants pollinated by wind

In such situations especially with some water plants mycoheterotrophs and

parasites a variety of characters may be affected and particularly under such

circumstances groups have tended to be formed using a few or even only a single

character that seems to provide evidence of relationships Add to this the tendency to

weight some characters particularly strongly on a priori grounds and the result was

either the recognition of conglomerate taxa such as Amentiferae which now appear to

be highly polyphyletic or the segregation of families like Plantaginaceae s str (now

much expanded) and Leitneriaceae (now in Simaroubaceae) - all of these are more or

less wind pollinated

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

18

Even although relationships of many parasitic and aquatic groups are being

clarified and at least some of the proposed phylogenetic positions seem to be well

supported it is indeed often still very difficult to understand how such groups have

evolved from their more morphologically conventional relatives However we have had

all sorts of preconceptions as to how characters should evolve Thus we used to

assume that features like highly scalariform vessel perforation plates or the complete

absence of vessels or a flower with an androecium that had many stamens a superior

ovary or separate petals were necessarily plesiomorphic or primitive and conversely

a simple perforation plate and the presence of vessels an androecium with few

stamens inferior ovary or petals that were connnate were almost necessarily

apomorphic or advanced (but cf eg Stebbins 1951) Such assumptions are incorrect

(eg Soltis et al 2005b) Carpels may become secondarily free carpels may fail to close

the seeds then developing outside the confines of the carpel as in some Aspagaraceae-

Nolinoideae Violaceae Berberidaceae Malvaceae-Sterculioideae etc) in Peliosanthes

teta perhaps the only species in Peliosanthes(Asparagaceae-Nolinoideae) the ovary

varies from superior to inferior (Jessop 1976 species limits here need close

investigation see also Kuzoff et al 2001 and Soltis amp Hufford 2002 Saxifragaceae

Apiales etc) many-seeded carpels can evolve from few-seeded carpels

(Razafimandimbison et al 2008) monoecy may be derived from dioecy (Schaefer amp

Renner 2010 and references) Classic studies such as those by Babcock (eg 1947)

on Crepis that assumed that evolution - in this case of the karyotype in particular - was

unidirectional have needed comprehensive re-evaluation (Enke amp Gemeinholzer 2008)

Most if not all characters have reversed andor evolved in parallel as is clear in the

discussion of ovary position here especially in Poales Saxifragales and Asparagales and

also in comments on such morphologically distinctive taxa (when compared with their

immediate relatives) such as Menyanthaceae and especially Pittosporaceae Parallel

evolution may occur even at the level of amino acid substitution as in the independent

acquisition of the phosphoenolpyruvate carboxylase (pepC) gene in C4 photosynthesis in

grasses (Christin et al 2007b see also Blaumlsing et al 2000)

Thinking of how such characters have been used in the past in classifications at the

generic level the results have been similar to those just discussed above for parasitic

plants and others Relying too much on animal pollination syndromes to mark generic

boundaries has all too often led to taxa that are highly unsatisfactory phylogenetically

(see eg Acanthaceae Bignoniaceae Campanulaceae Ericaceae Lamiaceae

Orchidaceae) and over-reliance on characters of fruit and seed (see eg Brassicaceae

and Apiaceae) has also led to unsatisfactory generic limits Again the more general

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

19

problem is the use of one or a very few characters that have been weighted a priori to

structure classifications (see also Garciacutea et al 2009) although it is perhaps not

surprising that such an approach sometimes results in the delineation of taxa that have

indeed turned out to be quite distinct (cf eg Burtt Astragalus versus Oxytropis)

Taxa recognized here are monophyletic yet many are polythetic at the

morphological level that is they lack unique features characterizing (defining) all and

only members of that family they can be recognised phenetically only by the unique

combinations of characters that they posess This is the result of evolution any

synapomorphy characterizing a taxon may be lost or modified beyond easy recognition

in some of its members or the synapomorphy may appear to be identical to a feature

that has evolved in parallel in a quite unrelated plant That plant groups are polythetic is

almost as much a feature of monophyletic taxa as the paraphyletic taxa common in

evolutionary classifications Some families now include substantial variation as

phenetically distinct derived groups are placed in their proper phylogenetic position -

examples are Ericaceae which include the erstwhile Empetraceae a wind-pollinated

group that is florally very different and the overwhelmingly large-flowered and animal-

pollinated Plantaginaceae which now include several derived small-flowered aquatic

and wind-pollinated groups that were previously placed in separate families

Thus some families as delimited here may not be easy to recognize However

remember that detecting relationships - use whatever characters you can even if they

are not obvious - and naming a plant - focus on easy-to-see characters that may not

reflect relationships - are quite different problems Taxa although natural may not be

readily recognizable indeed it was in exactly this context that Lamarck worked out the

basic principles of writing dichotomous keys in 1778 (see Scharf 2007 for keys and the

like) Of course Lamarcks idea of nature was very different from ours - he thought that

there was some kind of continuum of form on which living organisms were to be

situated with no real gaps anywhere - but this meant that his genera (for example)

might well not be sharply distinct from each other even if each were part of the real

continuum that was life Lamarcks separation of the establishment of relationships and

groupings in a system and identification of the plants in that system have been widely

accepted (but cf eg Godfray amp Knapp 2004 many of the contributors to Heywood et

al 2007) Perhaps the best way of identifying plants at the family level is by well-made

multiple access keys as in Watson and Dallwitz (1992a onwards family limits there may

differ substantially from those adopted here) Multiple access keys free users from the

constraints of dichotomous keys in which particular characters are needed at each step

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

20

of the identification process before there can be further progress Instead those

characters that happen to be evident on a specimen can be used in whatever order is

convenient when linked to illustrations glossaries etc their power is enormous (see

Dallwitz et al 2000 [2006] for the principles underlying their construction and use)

Nevertheless dichotomous keys such as those of Hutchinson (1973) and Franz Thonner

(Geesink et al 1981) have their uses Of course taxa coming out adjacent in keys may

well not be at all related

(When identifying large numbers of plants even more efficient than either style of

identification and certainly lots more fun is sight identification Unless you have a

photographic memory you have to build up your knowledge of comparative plant

morphology - on which the ability to make accurate identifications depends - by

repeated observation When faced with an unknown plant I always look for leaf teeth

and stipules distinctive hair types especially stellate and T-shaped hairs glands and

punctations of any sort the presence of latex or other exudate and check leaf insertion

smelling crushed leaves can also be helpful In this context nodal anatomy can usually

be checked using a razor and a hand lens or even simply carefully examining leaf scars

The short paragraphs added after most families may help in confirming familial

identifications)

For the record and for the little that it is worth there are 4 orders and 13 families

of gymnosperms characterised on these pages and together they include some 82

genera and 947 species For angiosperms comparable figures are 56 orders 445

families 13208 genera and 261750 species (of which monocots include 11 orders 89

families 2759 genera and 52760 species) Note however that higher mathematics

was never my strong point and anyway these are pretty meaningless figures even for

species which many (but not all) might concede smacked slightly more of reality than

other taxa estimates range as high as 422000 (Govaerts 2001) Furthermore numbers

of genera and species change daily Nevertheless as emphasized here families are

useful in teaching we as a community can ensure that their limits remain largely stable

and by concentrating on relatively few of them one can gain some familiarity with much

of the worlds flora For summaries of the sizes of orders and families and of the general

arrangement followed in these pages see also the Statistics page itself

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

21

A PHYLOGENETIC TREE OF SEED PLANTS (THE MAIN TREE)

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

22

The Main Tree above is a conservative summary of well supported relationships

between the major clades of extant seed plants Reading - interpreting and

understanding - such a phylogenetic tree is not necessarily easy so reading a tree may

require a bit of practice OHara (eg 1988 1992) provides an accessible introduction to

thinking and talking about history in the context of looking at phylogenetic trees More

recently Krell and Cranston (2004) Crisp and Cook (2005) and others have emphasized

how careful one must be when interpreting and talking about evolution in ladderized

trees in particular and phylogenetic trees in general Gregory (2008) provides a

particularly detailed discussion of this problem In ladderized trees the smaller (in terms

of numbers of terminals) sister taxon is consistently shown on the same side at every

node thus the trees tend to be pectinate like the teeth of a comb however the

horizontal axis that is as a result so evident has no polarity or particular direction The

use of the adjective basal when discussing phylogenetic trees is especially dangerous

(see also Wojciechowski et al 2004 D Soltis et al 2005b) When I use the term and the

context is not otherwise clear I am referring to the pectinations at the base of a

ladderised tree Remember that when talking about sister taxa one can never be basal

to the other or older or younger although members of one can be more derived - in the

sense of having more apomorphies - than the other (but even then one has to be

careful) The words primitive and advanced and lower and higher should

practically never be used especially when talking about taxa Amborellaceae and

Pinaceae are sister to all other flowering plants and Pinales and are basal to the crown

groups of all other flowering plantsPinales respectively but that does not mean they

are thereby primitive Pinaceae in particular have numerous apomorphies The word

plesiomorphic is far less loaded than primitive and can be used to talk about

individual characters

Most trees here have been more or less ladderized as is the one above The

ladderization may be imperfect for example you can see that on the Main Tree

asterids with ten orders follow rosids and relatives which have seventeen although

this is in part because of the currently unresolved nature of relationships between the

main clades of core eudicots (this is likely to change soon) The Main Tree could have

been drawn with Amborella Acorus or a host of other taxa at the far right without

offending any relationships phylogenetic trees are like mobiles the only fixed points

being the nodes Despite the problems just mentioned pectination interpreted

carefully has its value and it is not an arbitrary process As one reads the terminals of a

pectinate tree from left to right adjacent terminals will be separated by apomorphies

that will cumulate in an appropriate fashion When thinking of a book or a herbarium

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

23

sequence (see Haston et al 2007) this is of some value Nymphaeales and

Austrobaileyales are here adjacent on the tree but they could be separated by

hundreds of families in the sequence merely by rotating only the node from which

Nymphaeales and their sister group (all other angiosperms minus about 75 species)

arise If adjacent in a book or herbarium or as in these summary pages then it is

relatively easy to relate their characters both apomorphic and plesiomorphic but if

separated by hundreds of pages or two floors in a large building then it is less easy to

get anything from the sequence (Since all orders in the pages of the main part of this

site are preceded by the apomorphies of all nodes immediately below them in the seed

plant phylogeny and because of the linkages that have been built in to the site this

problem is much less serious there there is no sequence) So the particular sequence of

all seed plants that is based on a phylogenetic classification but is to be used in herbaria

or any other place where taxa have to be arranged linearly as in this part of APweb

will be one that maximizes the number of taxa that are both successive branches of the

tree and placed successively in the sequence Since specimens are generally filed under

families the outline of a new family sequence for arranging herbaria and books can now

be suggested (Haston et al 2007) and such a standardized sequence will further help

teaching and learning about plants

ON THINKING ABOUT APOMORPHIES

Identifying apomorphies is important because understanding the evolution of

morphology in the broad sense - ie including anatomy chemistry etc - in the context

of diversification is one of our major goals For identifying apomorphies several

preconditions must be met One needs to have an accurate well supported phylogeny

one has to have examined the right taxa both from the point of view of morphology and

molecules one has to have coded the characters right (ie delimited states

appropriately) and one has to use an appropriate model of evolution when fitting the

variation to the tree (see eg Omland 1999 Stevens 2006b) If the distributions of

apomorphies below are compared with those in other studies (eg Turgeon et al 2001

Bremer et al 2001 Endress 2001 Albach et al 2001a Judd amp Olmstead 2004 D Soltis

et al 2005b Zhang et al 2006 Judd et al 2007) the reader may find a number of

differences Although I have integrated such studies as far as is possible there are five

reasons that link to the preconditions just mentioned why there may be differences

where features are placed on trees

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

24

1 I may not have found all the information about a particular character there may be

disagreement over its interpretation or I have added information

2 The sampling of nearly all molecular studies is very incomplete (see Salisbury amp Kim

2001 for problems caused by sampling) indeed it is commonplace to decry the

incompleteness of molecular sampling But not only is the sampling in molecular studies

often less than we might wish that of the morphological and chemical characters whose

evolution we are interested in understanding is also often very poor So for many

anatomical chemical and embryological characters that are confidently said to

characterise families and other groups we all too often have no idea if those characters

are applicable to the whole clade or just to a subgroup within that clade However as

our sampling improves we can locate changes on the tree more precisely if in

unexpected positions Thus is is sometimes suggested that Ericaceae have ellagic acid

(Soltis et al 2005b) in fact the little evidence we have indicates that only a small clade

of some 80 species has ellagic acid and the rest of the family - itself alone about one

third of all Ericales - does not Similarly Ericaceae are noted for their

ectendomycorrhizal associations but Enkianthus sister to all other Ericaceae

apparently lacks these (Abe 2005) Thus these mycorrhizae and their associated

distinctive hair roots are unlikely to be apomorphies of Ericaceae - however it appears

that other families near Ericaceae may also have ectendomycorrhizae (see Asai 1934)

3 I am fitting characters to a very conservative tree with many polytomies although the

nodes that are utilised are for the most part strongly supported polytomies make the

optimisation of characters that is the assigment of character state change to a

particular node on the tree notably difficult (eg Madison amp Madison 2002) In nearly

all studies of the evolution of characters distributions of characters are optimised on a

more or less fully resolved tree and the construction of supertrees may yield yet more

detailed hypotheses of relationships (for literature on supertrees see Cotton amp

Wilkinson 2007 2008) Of course some nodes on such fully resolved trees andor

supertrees may have little support and optimisations of characters on such trees may

carry correspondingly little conviction Even parts of some of the trees used here have

poor support eg relationships within aquatic Alismatales etc although I indicate

when this is the case

4 Exactly how one goes about optimising a character on a tree is critically important

Even using simple parsimony optimisations (ACCTRAN or DELTRAN ACCelerated

TRANsitions or DELayed TRANsitions) the position of synapomorphies on trees - and

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

25

hence our ideas of evolution - may differ and this is still more true if one uses maximum

likelihood or Bayesian methods and makes apparently reasonable suggestions about

weighting gains over losses (eg Donoghue amp Ackerley 1996 Cunningham et al 1998

Omland 1997 1999 Ree amp Donoghue 1999 Polly 2001 Webster amp Purvis 2001

Ronquist 2004 Crisp amp Cook 2005) Sannier et al (2007) show how in in Arecaceae that

where on a phylogenetic tree one might peg changes in microsporogenesis will depend

on the methods one uses to do this and Pedersen et al (2007) discuss the sometimes

very substantial effect of node support on the posterior probabilities of ancestral

character states Here I use parsimony optimization not always as explicit as it might be

but I have often indicated where there are particularly important uncertainties as to the

positions of particular character changes on the tree

5 Finally although I have paid quite a lot of attention to the delimitation of the

character states that make up all the characterizations I have not spent enough time on

this critical operation If we are interested in understanding evolution then fitting the

basic variation - not character states - to a tree in principle allows greater flexibility in

understanding morphology in the context of local phylogenies (see also Stevens 2000

Endress 2005c) However many character states used here are delimited globally that

is they are circumscribed in the context of the variation shown by individual characters

across all angiosperms andor in the context of classic ideas of character evolution

Character states often have arbitrary limits and serve best to communicate

information whether they are in fact suitable for either phylogenetic analysis or

understanding evolution are separate issues Studies have rather unsurprisingly

perhaps but importantly shown that dividing the one character into different sets of

states may yield differing ideas of evolution of that character (eg Lamb Frye amp Kron

2003 Hibbett 2004) When looking at trees on which character states are optimised

one should bear in mind the problems surrounding the delimitation of states (eg

Stevens 2000 2006b) and the danger of using pollen or other types - constructs

based on many characters that vary independently but which effectively get lost in

these types thus Blackmore et al (2009) decomposed the pollen types in Asteraceae

into 52 characters

I use Remanes three main criteria of homology or better similarity when

determining the basic similarity of structures on different organisms (see Remane 1952)

These criteria are special properties position and intermediates Special properties

include anatomical or chemical characters gene expression data etc Position refers

to the position of an organ with respect to landmarks on the plant Although plants are

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

26

plants and landmarks are not as fixed as one might like trying to work out the relative

positions of parts is a good way of understanding morphology so long as one does not

try to out-do Procrustes Intermediates include intermediates found on the same

plant or on different plants Intermediates may observed during development or be

adult structures Thus one may find changes during development which suggest what

the nature of a particular structure is and very different stuctures on different plants

may turn out to be similar early in development Or one can simply compare different

taxa and see that structures that appear to be very different when comparing two

different taxa can be linked morphologically by looking at other taxa

Nevertheless the use of these criteria may not yield an unambiguous answer as to

what a structure is even given a solid phylogeny and an improved understanding of

development (see Jaramillo amp Kramer 2007 for a useful discussion) As Endress (2005c)

observed a number of features - position function development shape anatomy

histology gene activity and relationships to other taxa that clearly have petals - can be

used to distinguish a petal (for example) from other floral structures if a petal does not

have one of these features is it thereby not a petal Thus Maturen et al (2005) recently

found that floral organ diversity genes (B and C) were expressed in the large white

inflorescence bracts of Cornus (see also Costa et al 2005) Peney et al (2005) noted that

not all monosulcate pollen grains in monocots have the same developmental pathway

and that as a result such pollen might not have the same ancestral state Reeves and

Olmstead (2003) suggested that the genetic mechanisms causing monosymmetry in

Lamiales and Solanales were different and Serna and Martin (2006) described similar

problems with the development of hairs in Arabidopsis when compared with that of

hairs in Antirrhinum and Solanaceae Indeed as one perhaps might expect delimitation

of states and characters does not necessarily become easier with increasing knowledge

of development etc Thus Buzgo et al (2004) Matthews and Endress (2005) and others

have shown how hard it can be to distinguish between eg prophylls and other floral

structures as their behaviour is studied during the course of the development of the

flower Similarly at what concentration is a particular secondary metabolite deemed to

be present (Waterman (2007)

The validity of the approach used here that of fitting morphological variation to a

largely molecular-based tree may be questioned However I think it rather unlikely that

well-supported molecular branches will be overturned by morphological data Indeed

analyses of morphological data alone do provide support for many of the clades evident

in molecule-only analyses and analyses with morphological and molecular data

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

27

together may lead to increased support for clades (eg Hufford 1992 Nandi et al 1998

[but adding morphological data reduces support for a number of critical clades too]

Doyle amp Endress 2000 however in none of these papers is the use of morphology

without ambiguity) It is unfortunately clear that the use of morphology alone may not

suggest problems in the phylogenetic placement of taxa that later turn out to have been

wrongly included (eg Zhang et al 1992) Thus I have been wary of putting much weight

on clades that have only morphological support but note that I have treated molecule-

based clades with low bootstrap or jacknife support values (esp below 70) or low

posterior probabilies (below 095) likewise Although I may have been mistaken in

placing so much emphasis on molecular data in terms of providing the basic

phylogenetic framework for angiosperms morphological and molecular data are only

very rarely in irreconcilably strong conflict There are indeed a few places where the

conflict seems extreme These include the relative positions of the Monimiaceae and

Hernandiaceae (Laurales) the position of Hanguanaceae (Commelinales [as here] or

Zingiberales) and of Triplostegia (is it in Dipsacaceae or Valerianaceae - see

Dipsacales) Fossils are unlikely to affect the topologies of the trees presented here but

see below for their importance in understanding morphological evolution in general and

the evolution of angiosperms in particular Nevertheless some largely reject the idea

that trees based on molecular data alone can recover phylogenetic relationships

especially when branching points are old and prefer to used trees based on analysis of

morphological data including those taken from fossils (Hilton amp Bateman 2006 Farjon

2007)

All in all however the extent of the congruence between morphological and

molecular data is impressive and heartening and many clades can be characterised

morphologically It seemed in 1998 that there were no unambiguous morphological

synapomorphies for angiosperm orders (K Bremer 2000) and this is still true if by

unambiguous is meant non-homoplasious However many orders and other clades

have synapomorphies even if these may be indistinguishable at least at the current

level of morphological and developmental knowledge from parallel occurences

elsewhere As our morphological knowledge increases so too does the number of

apomorphies

Endress and Matthews (2006a) emphasize the importance of tendencies and

developmental constraints when thinking about characters of clades within the rosids -

many characters or character combinations occur in a rather sporadic fashion within

the clade and apparently notably less frequently outside it There are a number of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

28

examples of tendencies here and in the individual order pages such as the features

enclosed by parentheses in groups above the level of family eg see (cuticular waxes

as aggregated rodlets) for the commelinids and the discussion of the distribution of

polyandry that is flowers with numerous stamens in the asterid I + II groups etc

Tendencies may involve more than single characters When thinking of the

associations of features like integument thickness and vascularization of the integument

(Endress amp Matthews 2006a) one can imagine fairly simple developmental

preconditions being operative As Endress and Matthews (2006a) note it is difficult to

imagine vascular tissue developing in an integument only two cells thick Changes here

would be loosely correlated if morphologically linked However the strongly correlated

changes noted by Givnish et al (2005) are ecologically linked but are presumably

morphologicallydevelopmentally independent When there seem to be characters

evolving more or less together tests can be carried out to see if the changes are

concentrated on certain branches of the tree (eg Maddison 1990 Sanderson 1991

Maddison amp Maddison 2000) Such tests have rarely been carried out at higher levels in

angiosperms

One of the most striking examples of a tendency is the distribution of N-fixation

restricted as it is to a monophyletic group of four clades although it has arisen seven (or

perhaps several more) times independently within the N-fixing clade and several

members of two quite different kinds of bacteria are involved (eg Clawson et al 2004

Elliott et al 2007 Sprent amp James 2007) Other examples of work that bears on the issue

of tendencies include the findings that flowers of polysymmetrical Arabidopsis have

genes like TCP1 that are expressed asymmetrically during early development and TCP1

is a probable orthologue of the well-known CYC gene of Antirrhinum that is involved in

the development of monosymmetric slowers there (Cubas et al 2001 Costa et al 2005

etc) Parallelism might build on this underlying morphologically cryptic monosymmetry

even if details of the genetic mechanisms causing the monosymmetry evident in

particular groups may be different (Reeves amp Olmstead 2003 Cubas 2004 see above)

Similarly there have been several recent suggestions that the capability to synthesise a

particular metabolite may be switched off but not lost and so can sometimes be

reacquired (eg Wink amp Witte 1983 Wink 2003 Liscombe et al 2005 Larsson 2007

Waterman 2007) Hence perhaps the rather spotty distribution of many secondary

metabolites like ellagic acid the indole alkaloid camptothecin iridoids etc when

considered in the context of phylogenies As a non-botanical example - but a rather nice

one - Salwini-Plawen and Mayr (1961) suggested some time ago that there has been

considerable parallelism (40-65 or more independent origins) in the evolution of eyes in

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

29

metazoans However Pax 6 seems to be a master control gene that is involved in eye

formation perhaps throughout all bilateralians (eg Gehring amp Ikeo 1999 Erwin amp

Davidson 2002) which changes the nature of the problem Note however that in

general our invocation of evolutionary tendencies to explain the patterns of

distributions of characters is really an admission that we do not understand those

patterns

As we find out more about variation we will probably find fewer and fewer features

constant throughout a group Most unqualified statements of presence and absence

should properly be qualified as usually present or usually absent if one is thinking of

the characterisations on the order pages as encompassing the total variation within a

clade Thus Pistia alone among monocots as so far known has sieve tube plastids with

starch grains not protein crystals However this does not affect the fact that sieve tube

plastids with cuneate protein crystals and lacking starch grains are an apomorphy for

monocots Furthermore fossils in a number of cases suggest character combinations

unknown in extant taxa as may be seen in the discussions of Fagaceae Platanaceae

Iteaceae Calycanthaceae etc Confusing the issue there may be questions as to where

exactly on the tree a particular fossil is to be placed (eg see Nymphaeaceae

Calycanthaceae Archaefructus etc)

To summarize given our current understandings of both phylogenies and

characters evolution of some characters in which we are interested seems very labile

(see eg D Soltis et al 2005b Endress and Matthews 2006a Stevens 2006b Ekman et

al 2008 for a good recent study) and I have been cautious when talking about character

evolution Much effort must continue to be spent in summarizing characters of well-

established clades at all levels providing features by which they may be recognized and

signaling synapomorphies Remember that (1) the basic morphological anatomical and

chemical knowledge of many critical taxa is woefully incomplete (2) different

assumptions about character evolution may greatly affect the position of

synapomorphies on trees (3) in many cases relationships within and between many

groups are too uncertain at present to worry very much about synapomorphies and

(4) we must be clear about what we do and do not not know As mentioned above I

have indicated in a number of places where there are particularly important

uncertainties as to where characters should be placed on the tree Nevertheless it is a

relatively easy matter to update notes such as these and it can be a simple matter to

incorporate new data on characters that have never before been considered in the

context of a tree Much basic - and unfortunately perhaps unfashionable - work must be

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

30

carried to clarify the distribution of morphological anatomical and chemical characters

for examples of what can to be done see the work of P K Endress and collaborators

(floral morphology and development) and S R Jensen and collaborators (iridoids) But

acquiring information about nodal anatomy cell and tissue distribution nodal anatomy

and the like is not difficult at all and should be generally encouraged

But all this is merely a necesary prelude to the understanding of evolution For this

one needs to know a lot more including dating the phylogeny understanding the

function(s) of characters factoring in relevant aspects of the palaeoenvironment in

which an apomorphy first appeared etc

SUMMARY OF THE SYSTEM

Below is a formalised summary of the relationships within orders of the families of seed

plants There are a few families that are not recognised even as options in APG II and

vice versa as well as a few extra orders The families to be recognised in the most

recent edition of Mabberleys The Plant Book (Mabberley 2008) are also largely

consistent with those below But all differences are trivial and will - I hope - eventually

disappear

Square brackets - [] - enclose clades the plus sign - + - designates sister taxa a

comma - - denotes part of a polytomy and quotation marks - - denotes a

paraphyletic group

SEED PLANTS

GYMNOSPERMS

Cycadales

Cycadaceae + Zamiaceae

Ginkgoales

Ginkgoaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

31

Pinales

Pinaceae [[Araucariaceae [Podocarpaceae + Phyllocladaceae]] [Sciadopityaceae

[Taxaceae + Cupressaceae]]]

Gnetales

Ephedraceae [Gnetaceae + Welwitschiaceae]

ANGIOSPERMSFLOWERING PLANTS

Amborellales

Amborellaceae

Nymphaeales

Hydatellaceae [Cabombaceae + Nymphaeaceae]

Austrobaileyales

Austrobaileyaceae [Schisandraceae + Trimeniaceae]

Chloranthales

Chloranthaceae

MAGNOLIIDS

Magnoliales

Myristicaceae [Magnoliaceae [[Himantandraceae + Degeneriaceae] [Eupomatiaceae +

Annonaceae]]]

Laurales

Calycanthaceae [[Siparunaceae [Gomortegaceae + Atherospermataceae]] [Monimiaceae

[Hernandiaceae + Lauraceae]]]

Canellales

Canellaceae + Winteraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

32

Piperales

[Hydnoraceae + Aristolochiaceae] [Piperaceae + Saururaceae]

MONOCOTS

Acorales

Acoraceae

Alismatales

Araceae [Tofieldiaceae [[Alismataceae [Hydrocharitaceae + Butomaceae]]

[Scheuchzeriaceae [Aponogetonaceae [Juncaginaceae [Maundiaceae [[Posidoniaceae

[Ruppiaceae + Cymodoceaceae]] [Zosteraceae + Potamogetonaceae]]]]]]]

Petrosaviales

Petrosaviaceae

Dioscoreales

Nartheciaceae [[Taccaceae + Thismiaceae] [Burmanniaceae + Dioscoreaceae]]

Pandanales

Velloziaceae Triuridaceae Stemonaceae [Pandanaceae + Cyclanthaceae]

Liliales

Corsiaceae [Campynemataceae [Petermanniaceae [Colchicaceae + Alstroemeriaceae]]

Melanthiaceae [[Philesiaceae + Rhipogonaceae] [Smilacaceae + Liliaceae]]]

Asparagales

Orchidaceae [[Boryaceae [Blandfordiaceae [Lanariaceae [Asteliaceae + Hypoxidaceae]]]]

[[Ixioliriaceae + Tecophilaeaceae] [Doryanthaceae [Iridaceae [Xeronemataceae

[Xanthorrhoeaceae [Alliaceae + Asparagaceae]]]]]]]

COMMELINIDS

Unplaced

Dasypogonaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

33

Arecales

Arecaceae

Poales

[Typhaceae + Bromeliaceae] [Rapateaceae [[Thurniaceae [Juncaceae + Cyperaceae]]

[[Mayacaceae [Eriocaulaceae + Xyridaceae]] [Flagellariaceae [Anarthriaceae

[Centrolepidaceae + Restionaceae]]] [Joinvilleaceae [Ecdeiocoleaceae + Poaceae]]]]]]]

Commelinales

[Commelinaceae + Hanguanaceae] [Philydraceae [Haemodoraceae + Pontederiaceae]]

Zingiberales

Musaceae [Strelitziaceae + Lowiaceae] Heliconiaceae [[Cannaceae + Marantaceae]

[Costaceae + Zingiberaceae]]

Ceratophyllales

Ceratophyllaceae

EUDICOTS

Ranunculales

Eupteleaceae [Papaveraceae [[[Lardizabalaceae + Circaeasteraceae] [Menispermaceae

[Berberidaceae + Ranunculaceae]]]]

Sabiales

Sabiaceae

Proteales

Nelumbonaceae [Platanaceae + Proteaceae]

Trochodendrales

Trochodendraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

34

Buxales

Haptanthaceae [Buxaceae + Didymelaceae]

CORE EUDICOTS

Gunnerales

Gunneraceae + Myrothamnaceae

Dilleniales

Dilleniaceae

Saxifragales

Peridiscaceae [[Paeoniaceae [Altingiaceae [Hamamelidaceae [Cercidiphyllaceae +

Daphniphyllaceae]]]] [[Crassulaceae [Aphanopetalaceae [Tetracarpaeaceae

[Penthoraceae + Haloragaceae]]]] [Iteaceae [Grossulariaceae + Saxifragaceae]]]]

Cynomoriaceae unplaced

Vitales

Vitaceae

ROSIDS

FABIDROSID I

Zygophyllales

Krameriaceae + Zygophyllaceae

Celastrales

Lepidobotryaceae + Celastraceae

Oxalidales

Huaceae [[Connaraceae + Oxalidaceae] [Cunoniaceae [Elaeocarpaceae [Brunelliaceae +

Cephalotaceae]]]]

Malpighiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

35

[Achariaceae [Goupiaceae [Violaceae + Passifloraceae] [Lacistemataceae Salicaceae]]

[Lophopyxidaceae Putranjivaceae] [Ctenolophonaceae [Erythroxylaceae +

Rhizophoraceae]] Linaceae Ixonanthaceae Humiriaceae Irvingiaceae

Centroplacaceae Pandaceae [[Bonnetiaceae + Clusiaceae] [Calophyllaceae

[Hypericaceae + Podostemaceae]]] [Malpighiaceae + Elatinaceae] Ochnaceae

[Peraceae [Rafflesiaceae + Euphorbiaceae]] [Phyllanthaceae + Picrodendraceae]

[Balanopaceae [[Trigoniaceae + Dichapetalaceae] [Chrysobalanaceae +

Euphroniaceae]]] Caryocaraceae

N-FIXING CLADE

Fabales

Quillajaceae [Fabaceae [Polygalaceae + Surianaceae]]

Rosales

Rosaceae [[Barbeyaceae [Elaeagnaceae Dirachmaceae Rhamnaceae]] [Ulmaceae

[Cannabaceae [Moraceae + Urticaceae]]]]

Cucurbitales

Anisophylleaceae [[Corynocarpaceae + Coriariaceae] [Cucurbitaceae [Tetramelaceae

[Datiscaceae + Begoniaceae]]]] Apodanthaceae

Fagales

Nothofagaceae [Fagaceae [[Myricaceae + Juglandaceae] [Casuarinaceae

[Ticodendraceae + Betulaceae]]]]

MALVIDROSID II

pgtGeraniales

Geraniaceae [[Melianthaceae + Francoaceae] [Vivianaceae + Ledocarpaceae]]

Myrtales

Combretaceae [[Onagraceae + Lythraceae] [[Vochysiaceae + Myrtaceae]

[Melastomataceae [Crypteroniaceae [Alzateaceae + Penaeaceae]]]]]

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

36

Crossosomatales

[Staphyleaceae [Guamatelaceae [Crossosomataceae + Stachyuraceae]]] [Aphloiaceae

[Geissolomataceae + Strasburgeriaceae]]

Picramniales

Picramniaceae

Sapindales

Biebersteiniaceae Nitrariaceae [Kirkiaceae [Anacardiaceae + Burseraceae]]

[Sapindaceae [Simaroubaceae Rutaceae Meliaceae]]

Huerteales

Gerradinaceae [Dipentodontaceae + Tapisciaceae]

Malvales

Neuradaceae [Thymelaeaceae [Sphaerosepalaceae Bixaceae [Cistaceae

[Sarcolaenaceae + Dipterocarpaceae]] [Cytinaceae + Muntingiaceae] Malvaceae]]

Brassicales

[Akaniaceae + Tropaeolaceae] [[Moringaceae + Caricaceae] [Setchellanthaceae

[Limnanthaceae [[Koeberliniaceae [Bataceae + Salvadoraceae]] [Emblingiaceae

[Pentadiplandraceae [Gyrostemonaceae Resedaceae Stixis and relatives] Tovariaceae

[Capparaceae [Cleomaceae + Brassicaceae]]]]]]]]

Berberidopsidales

Aextoxicaceae + Berberidopsidaceae

Santalales

Erythropalaceae Olacaceae [[Loranthaceae [Misodendraceae + Schoepfiaceae]]

[Opiliaceae + Santalaceae]] Balanophoraceae unplaced

Caryophyllales

[[Droseraceae [Nepenthaceae [Drosophyllaceae [Ancistrocladaceae +

Dioncophylleaceae]]]]] [[Frankeniaceae + Tamaricaceae] [Polygonaceae +

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

37

Plumbaginaceae]]] [Rhabdodendraceae [Simmondsiaceae [[Asteropeiaceae +

Physenaceae] [[Caryophyllaceae [Achatocarpaceae + Amaranthaceae]]

[Stegnospermataceae [Limeaceae [[Lophocarpaceae [Barbeuiaceae [Aizoaceae

[Sarcobataceae Phytolaccaceae Nyctaginaceae]]]] [Molluginaceae [Halophytaceae

Montiaceae Basellaceae Didiereaceae [Talinaceae [Portulacaceae [Anacampseros etc +

Cactaceae]]]]]]]]]]]]]

ASTERIDS

Cornales

[Cornaceae + Nyssaceae] [Hydrangeaceae + Loasaceae] Hydrostachyaceae [Curtisiaceae

+ Grubbiaceae]

Ericales

[Balsaminaceae [Marcgraviaceae + Tetrameristaceae]] [[Polemoniaceae +

Fouquieraceae] Lecythidaceae [[Sladeniaceae + Pentaphylacaceae] [Sapotaceae

[Ebenaceae [Maesaceae [Theophrastaceae [Myrsinaceae + Primulaceae]]]]]

[Mitrastemonaceae Theaceae [Symplocaceae [Styracaceae + Diapensiaceae]]

[[Sarraceniaceae [Roridulaceae + Actinidiaceae] [Clethraceae [Cyrillaceae +

Ericaceae]]]]]]

LAMIIDASTERID I

Unplaced

Metteniusaceae Oncothecaceae Icacinaceae and some unplaced ex-Icacinaceae

Garryales

Garryaceae + Eucommiaceae

Unplaced

Boraginaceae Vahliaceae

Gentianales

Rubiaceae [Gentianaceae [Loganiaceae [Gelsemiaceae + Apocynaceae]]]

Lamiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

38

Plocospermataceae [Oleaceae [Tetrachondraceae [Calceolariaceae

[Gesneriaceae Peltanthera [Pedaliaceae Carlemanniaceae Martyniaceae Stilbaceae

Plantaginaceae Linderniaceae Bignoniaceae Verbenaceae Lamiaceae Paulowniaceae

Schlegeliaceae Thomandersiaceae Phrymaceae Rehmannia Acanthaceae

Scrophulariaceae Orobanchaceae Byblidaceae Lentibulariaceae]]]]]

Solanales

[Montiniaceae [Sphenocleaceae + Hydroleaceae]] [Convolvulaceae + Solanaceae]

CAMPANULIDASTERID II

Aquifoliales

[[Cardiopteridaceae + Stemonuraceae] [Aquifoliaceae [Helwingiaceae +

Phyllonomaceae]]

Asterales

[Rousseaceae + Campanulaceae] Pentaphragmataceae [[Alseuosmiaceae [Phellinaceae

+ Argophyllaceae]] Stylidiaceae [Menyanthaceae [Goodeniaceae [Calyceraceae +

Asteraceae]]]]

Escalloniales

Escalloniaceae

Bruniales

[Bruniaceae + Columelliaceae]

Apiales

Pennantiaceae [Torricelliaceae [Griseliniaceae [Pittosporaceae [Araliaceae

[Myodocarpaceae + Apiaceae]]]]]

Paracryphiales

Paracryphiaceae

Dipsacales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

39

Adoxaceae [Diervillaceae [Caprifoliaceae [Linnaeaceae [Morinaceae [Dipsacaceae +

Valerianaceae]]]]]

LINKS TO ORDERS AND FAMILIES

[Back to Top]

MAIN GROUPINGS

Asterids commelinids core eudicots asterid 1 asterid

2 eudicots gymnosperms Magnoliophyta monocots N-fixing claderosids seed plants

ALPHABETICAL LISTING OF ALL ORDINAL NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | L | M | N | O | P | Q | R | S | T | U | V | W | X | Z

Abietales Acanthales Acerales Acorales Actinidiales Actinostrobales Adoxales Aescul

ales Agavales Aizoales AkanialesAlismatales Alliales Alseuosmiales Alstroemeriales

Altingiales Amaranthales Amaryllidales Amborellales AmbrosialesAmmiales Amomal

es Ancistrocladales Anisophylleales Annonales Anthobolales Apiales Apocynales Apo

nogetonalesAquifoliales Arales Araliales Aralidiales Araucariales Arecales Aristolochi

ales Athrotaxidales Asarales AsclepiadalesAsparagales Asphodelales Asparagales Ast

eliales Atriplicales Aucubales Austrobaileyales Avenales

Balanitales Balanopales Balanophorales Balsaminales Barbeyales Barclayales Batales

Begoniales BerberidalesBerberidopsidales Betulales Biebersteiniales Bignoniales Bix

ales Boraginales Brassicales Brexiales Bromeliales BrunialesBrunoniales Burmannial

es Burserales Butomales Buxales Byblidales

Cactales Callitrichales Calycanthales Calycerales Campanulales Campynematales Can

ellales Cannales CapparalesCaprifoliales Cardiopteridales Carduales Caricales Carle

manniales Caryophyllales Cassiales Casuarinales CelastralesCentrolepidales Cephalot

ales Cephalotaxales Ceratophyllales Cercidiphyllales Chenopodiales Chironiales Chlor

anthalesChrysobalanales Cinchonales Circaeasterales Cistales Citrales Cocosales Col

chicales Columelliales CombretalesCommelinales Connarales Convolvulales Coriarial

es Cornales Corylales Corynocarpales Crassulales CrossosomatalesCucurbitales Cunn

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

40

inghamiales Cunoniales Cupressales Cycadales Cyclanthales Cymodoceales Cynarales

CynomorialesCyperales Cyrillales Cytinales

Daphnales Daphniphyllales Dasypogonales Datiscales Degeneriales Desfontainiales D

iapensiales Didymelales DillenialesDioales Dioncophyllales Dioscoreales Diospyrales

Dipentodontales Dipsacales Droserales

Ebenales Echiales Elaeagnales Elaeocarpales Elatinales Elodeales Emmotales Empetr

ales Ephedrales Ericales EriocaulalesErythropalaless Escalloniales Eucommiales Euph

orbiales Eupomatiales Eupteleales Euryalaless

Fabales Fagales Falcatifoliales Ficales Flacourtiales Flagellariales Fouquieriales Franc

oales Frangulales

Galiales Garryales Geissolomatales Gentianales Geraniales Gesneriales Ginkgoales G

laucidiales Globulariales GnetalesGoodeniales Greyiales Griseliniales Grossulariales

Grubbiales Gunnerales Gyrocarpales Gyrostemonales

Haemodorales Haloragales Hamamelidales Hanguanales Heisteriales Helleborales He

lwingiales Himantandrales HippuridalesHomaliales Hortensiales Huales Huerteales

Hydatellales Hydnorales Hydrangeales Hydrastidales HydrocharitalesHydropeltidales

Hydrostachyales Hypericales Hypoxidales

Icacinales Illiciales Iridales Irvingales Iteales Ixiales

Jasminales Juglandales Julianiales Juncaginales Juncales

Lacistematales Lactoridales Lamiales Lardizabalales Laurales Lecythidales Ledocarpal

es Leitneriales LentibularialesLigustrales Liliales Limnanthales Linales Liriales Loasal

es Lobeliales Loganiales Lonicerales Loranthales LowialesLythrales

Magnoliales Malpighiales Malvales Marathrales Marcgraviales Mayacales Medusagy

nales Medusandrales MelanthialesMelastomatales Meliales Melianthales Meliosmal

es Menispermales Menyanthales Metteniusales MitrastemonalesMiyoshiales Moni

miales Moringales Musales Myricales Myristicales Myrothamnales Myrsinales Myrta

les

Najadales Nandinales Narcissales Nartheciales Nelumbonales Nepenthales Neuradal

es Nitrariales Nolanales NothofagalesNyctaginales Nymphaeales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

41

Ochnales Oenotherales Olacales Oleales Onagrales Oncothecales Opuntiales Orchid

ales Oxalidales

Saxifragales Pandales Pandanales Papaverales Paracryphiales Parasitaxales Paridales

Parnassiales PassifloralesPenaeales Pennantiales Pentaphragmatales Peridiscales P

etiveriales Petrosaviales Phellinales Philydrales PhyllanthalesPhysenales Phytolaccal

es Picramniales Pinales Pinguiculales Piperales Pittosporales Plantaginales Platanale

sPlumbaginales Poales Podocarpales Podophyllales Podostemales Polemoniales Pol

ygalales Polygonales PontederialesPortulacales Posidoniales Potamogetonales Primu

lales Proteales

Quercales Quillajales Quintiniales

Rafflesiales Ranunculales Rapateales Resedales Restionales Rhabdodendrales Rhamn

ales Rhinanthales RhizophoralesRhodorales Rhoipteleales Roridulales Rosales Rouss

eales Rubiales Ruppiales Rutales

Sabiales Salicales Salvadorales Samolales Samydales Sanguisorbales Santalales Sapin

dales Sapotales SarracenialesSaxegotheales Saxifragales Scheuchzeriales Sciadopitya

les Scleranthales Scrophulariales Scyphostegiales SedalesSilenales Simmondsiales S

milacales Solanales Sphenocleales Sphenostemonales Stangeriales Stellariales Stemo

nalesStilbales Stylidiales Styracales Surianales

Taccales Tamales Tamaricales Taxales Taxodiales Tecophilaeales Terebinthales Tern

stroemiales Theales TheligonalesThymelaeales Tiliales Tofieldiales Torricelliales Tov

ariales Tribelales Trilliales Trimeniales Triuridales TrochodendralesTropaeolales Turn

erales Typhales

Ulmales Urticales

Vacciniales Vahliales Vallisneriales Velloziales Veratrales Verbenales Viburnales Vinc

ales Violales Viscales VitalesVochysiales

Welwitschiales Winterales

Xanthorrhoeales Ximeniales Xyridales

Zamiales Zingiberales Zosterales Zygophyllales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

42

ALPHABETICAL LISTING OF ALL FAMILY NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | K | L | M | N | O | P | Q | R | S | T | U | V | W | X | Y

|Z

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which the genus of the basionym is to be found

Abietaceae Abolbodaceae Abrophyllaceae Acalyphaceae Acanthaceae [Acanthochla

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ranthaceae Acmopylaceae Acoraceae Acristaceae ActaeaceaeActinidiaceae Actinota

ceae Amentotaxaceae Adenogrammaceae Adoxaceae Aegialitidaceae Aegicerataceae

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eae Agavaceae AgdestidaceaeAgialidaceae Agrimoniaceae Agrostidaceae Ailanthace

ae Aitoniaceae Aizoaceae Akaniaceae Alangiaceae AlchemillaceaeAldrovandaceae A

lismataceae Alliaceae Allioniaceae Allophylaceae Aloaceae Alopecuraceae Alpiniacea

e AlseuosmiaceaeAlsinaceae Alsodeiaceae Alstroemeriaceae Altingiaceae Alzateacea

e Amaranthaceae Amaryllidaceae Amborellaceae Ambrosiaceae Amentotaxaceae A

mmanniaceae Ammiaceae Amomaceae Ampelidopsaceae Amygdalaceae Amyridacea

eAnacardiaceae Anagallidaceae Anarthriaceae Anchusaceae Ancistrocladaceae Andr

omedaceae AndropogonaceaeAndrostachyaceae Androsynaceae Anemarrhenaceae

Anemonaceae Angelicaceae Anisophylleaceae AnnonaceaeAnomochloaceae Anopter

aceae Anrederaceae Anthemidaceae Anthericaceae Anthobolaceae Antidesmataceae

Antirrhinaceae Antoniaceae Aparinaceae Aphanopetalaceae Aphloiaceae Aphyllanth

aceae Apiaceae ApocynaceaeApodanthaceae Aponogetonaceae Aporusaceae Apose

ridaceae Apostasiaceae Aptandraceae AquifoliaceaeAquilariaceae Aquilegiaceae A

raceae Aragoaceae Araliaceae Aralidiaceaee Araucariaceae Arbutaceae Arceuthidace

aeArceuthobiaceae Arctostaphylaceae Arctotidaceae Ardisiaceae Arecaceae Argoph

yllaceae Arisaraceae AristolochiaceaeAristoteliaceae Arjonaceae Armeriaceae Artem

isiaceae Arthrotaxidaceae Artocarpaceae Arundinaceae ArundinellaceaeAsaraceae A

sclepiadaceae Ascyraceaee Asiraceae Asparagaceae Asperulaceae Asphodelaceae As

pidistraceaeeAsteliaceae Asteraceae Asteranthaceae Asteropeiaceae Astragalaceae

Astrocarpaceae AthanasiaceaeAtherospermataceae Atriplicaceae Atropaceae Aucuba

ceae Aurantiaceae Austrobaileyaceae Austrotaxaceae AvenaceaeAverrhoaceae Avet

raceae Avicenniaceae Azaleaceae Azimaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Balanitaceae Balanopaceae Balanophoraceae Balsameaceae Balsaminaceae Bambusa

ceae Barbaceniaceae BarbeuiaceaeBarbeyaceae Barclayaceae Barringtoniaceae Base

llaceae Bataceae Baueraceae Bauhiniaceae Baxteriaceae BegoniaceaeBehniaceae B

elangeraceae Belloniaceae Bembiciaceae Berberidaceae Berberidopsidaceae Berryac

eaee BersamaceaeBertyaceae Berzeliaceae Besleriaceae Betaceae Betulaceae Biebe

rsteiniaceae Bifariaceae Bignoniaceae BischofiaceaeBixaceae Blakeaceae Blandfordia

ceae Blattiaceae Blepharocaryaceae Blitaceae Blyxaceae Boerlagellaceae Bolivaracea

eBombacaceae Bonnetiaceae Bontiaceae Boopidaceae Boraginaceae Borassaceae B

oroniaceae BoryaceaeBotryodendraceae Bougainvilleaceae Boweniaceae Brassicacea

e Brachycaulaceae Bretschneideraceae BrexiaceaeBromeliaceae Brownlowiaceae Br

unelliaceae Bruniaceae Brunoniaceae Brunsvigiaceae Bryoniaceae BuchneraceaeBuci

daceae Buddlejaceae Buglossaceae Bulbocodiaceae Bumeliaceae Bupleuraceae Burc

hardiaceae BurmanniaceaeBurseraceae Butneriaceae Butomaceae Buxaceae Byblida

ceae Byttneriaceae

Cabombaceae Cacaoaceae Cactaceae Caesalpiniaceae Caladiaceae Calamaceae Calce

olariaceae CalectasiaceaeCalendulaceae Callaceae Callicomaceae Calligonaceae Calli

traceae Callitrichaceae Calochortaceae CalophyllaceaeCalthaceae Calycanthaceae Ca

lyceraceae Cambogiaceae Camelliaceae Campanulaceae CampynemataceaeCanacom

yricaceae Canellaceae Cannabaceae Cannaceae Canopodaceae Canotiaceae Cansjera

ceae CapparaceaeCaprariaceae Caprifoliaceae Cardiopteridaceae Carduaceae Carica

ceae Carissaceae Carlemanniaceae CarpinaceaeCarpodetaceae Cartonemataceae Ca

ryocaraceae Caryophyllaceae Casearieaceae Cassiaceae CassipoureaceaeCassythacea

e Castaneaceae Castelaceae Casuarinaceae Catesbaeaceae Cathedraceae Cecropiace

ae CedraceaeCedrelaceae Celastraceae Celosiaceae Celtidaceae Centaureaceae Cen

trolepidaceae Centroplacaceae CepaceaeCephalanthaceae Cephalotaceae Cephalota

xaceae Cerastiaceae Ceratoniaceae Ceratophyllaceae CerberaceaeCercidiphyllaceae

Cercocarpaceae Cercodiaceae Cereaceae Cerinthaceae Ceroxylaceae Cestraceae Cev

alliaceaeChailletiaceae Chamaedoraceae Chamaemoraceae Chamelauciaceae Chaun

ochitonaceae Chelidoniaceae ChelonaceaeChenopodiaceae Chimonanthaceae Chingi

thamnaceae Chionographidaceae Chiranthodendraceae ChironiaceaeChlaenaceae Ch

loanthaceae Chloranthaceae Chloridaceae Chlorogalaceae Chrysobalanaceae Cicerac

eae CichoriaceaeCimicifugaceae Cinchoniaceae Circaeaceae Circaeasteraceae Cissac

eae Cistaceae Citraceae Clematidaceae CleomaceaeClethraceae Clusiaceae Cneorac

eae Cnestidaceae Cnicaceae Cobaeaceae Cochlospermaceae Cocosaceae Coffeaceae

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Colchicaceae Coleogynaceae Columelliaceae Combretaceae Commelinaceae Comocl

adiaceae Compositae CompsoaceaeConantheraceae Connaraceae Conostylidaceae C

onvallariaceae Convolvulaceae Cordiaceae CoreopsidaceaeCoriandraceae Coriariacea

e Coridaceae Corispermaceae Cornaceae Corokiaceae Coronillaceae Correaceae Cor

rigiolaceaeCorsiaceae Corydalaceae Corylaceae Corynocarpaceae Coryphaceae Cost

aceae Cotyledonaceae CoulaceaeCoutariaceae Coutoubiaceae Crassulaceae Crescen

tiaceae Cressaceae Crinaceae Crocaceae CroomiaceaeCrossosomataceae Crotonacea

e Cruciferae Cryptaceae Crypteroniaceae Cryptocorynaceae CryptomeriaceaeCtenol

ophonaceae Cucurbitaceae Cunninghamiaceae Cunoniaceae Cupressaceae Curcumac

eae Curtisiaceae CuscutaceaeCyananthaceae Cyanastraceae Cynanchaceae Cyanella

ceae Cycadaceae Cyclanthaceae CyclantheraceaeCyclocheilaceae Cydoniaceae Cymo

doceaceae Cynaraceae Cynocrambaceae Cynomoriaceae Cyperaceae CyphiaceaeCyp

hocarpaceae Cypripediaceae Cyrillaceae Cyrtandraceae Cyrtanthaceae Cytinaceae

Dacrycarpaceae Dactylanthaceae Damasoniaceae Daphnaceae Daphniphyllaceae Das

ypogonaceae DatiscaceaeDaturaceae Daucaceae Davidiaceae Davidsoniaceae Decais

neaceae Deeringiaceae Degeneriaceae DelphiniaceaeDendrophthoaceae Columelliac

eae Detariaceae Dialypetalanthaceae Dianellaceae Dianthaceae DiapensiaceaeDicha

petalaceae Dichondraceae Diclidantheraceae Dicrastylidiaceae Dictamnaceae Didiere

aceae DidymelaceaeDidymocarpaceae Diegodendraceae Diervillaceae Digitalidaceae

e Dilatridaceae Dilleniaceae Dioaceae DionaeaceaeDioncophyllaceae Dioscoreaceae

Diosmaceae Diospyraceae Dipentodontaceae Diphylleiaceae Diplolaenaceae Dipsaca

ceaeDipterocarpaceae Dirachmaceae Disanthaceae Diselmaceae Dodonaeaceae Do

mbeyaceae Donatiaceae DorsteniaceaeDortmannaceae Doryanthaceae Drabaceae D

racaenaceae Dracontiaceae Drimyidaceae Droseraceae DrosophyllaceaeDryadaceae

Duabangaceae Duckeodendraceae Dulongiaceae Durantaceae Durionaceae Dysphani

aceae

Ebenaceae Eccremidaceae Ecdeiocoleaceae Echiaceae Echinopaceae Ehretiaceae Ela

eagnaceae ElaeocarpaceaeElegiaceae Elatinaceae Ellisiaceae Ellisiophyllaceae Elodea

ceae Elytranthaceae Embeliaceae Emblingiaceae EmmotaceaeEmpetraceae Encepha

lartaceae Engelhardtiaceae Enhalaceae Epacridaceae Ephedraceae Ephemeranthacea

eEpilobiaceae Epimediaceae Eragrostidaceae Eremolepidaceae Eremosynaceae Erica

ceae Erinaceae EriocaulaceaeEriogonaceae Eriospermaceae Erodiaceae Erycibaceae

Eryngiaceae Erysimaceae Erythroniaceae ErythropalaceaeErythrospermaceae Erythro

xylaceae Escalloniaceae Eschscholziaceae Eucomidaceae Eucommiaceae Eucryphiace

aeEuonymaceae Eupatoriaceae Euphorbiaceae Euphrasiaceae Euphroniaceae Eupo

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matiaceae Eupteleaceae EuryalaceaeEustrephaceae Euthemidaceae Eutocaceae Exb

ucklandiaceae Exocarpaceae

Fabaceae Fagaceae Falcatifoliaceae Ferulaceae Festucaceae Ficaceae Ficoidaceae Fit

zroyaceae FlacourtiaceaeFlagellariaceae Flindersiaceae Foetidiaceae Forestieraceae

Fothergillaceae Fouquieriaceae Fragariaceae FrancoaceaeFrangulaceae Frankeniacea

e Fraxinaceae Fraxinellaceae Freycinetiaceae Fritillariaceae Fuchsiaceae Fumariacea

eFunkiaceae

Gaiadendraceae Galacaceae Galanthaceae Galaxiaceae Galedupaceae Galeniaceae G

aliaceae Garciniaceae GardeniaceaeGardneriaceae Garryaceae Geissolomataceae Ge

itonoplesiaceae Gelsemiaceae Geniostomaceae GentianaceaeGeonomataceae Geosir

idaceae Geraniaceae Gerrardinaceae Gesneriaceae Gethyllidaceae Gilliesiaceae Gina

lloaceaeGinkgoaceae Gisekiaceae Gladiolaceae Glaucidiaceae Glechomaceae Glinace

ae Globulariaceae Gnaphaliaceae GnetaceaeGoetzeaceae Gomortegaceae Gomphiac

eae Gomphrenaceae Gonystylaceae Goodeniaceae Gordoniaceae GouaniaceaeGoup

iaceae Gramineae Gratiolaceae Grewiaceae Greyiaceae Grielaceae Griseliniaceae Gr

onoviaceae GrossulariaceaeGrubbiaceae Guaiacanaceae Guamatelaceae Guettardac

eae Gunneraceae Gustaviaceae Guttiferae GyrocarpaceaeGyrostemonaceae

Hachetteaceae Haemanthaceae Haemodoraceae Halesiaceae Hallieraceae Halocarpa

ceae Halophilaceae HalophytaceaeHaloragaceae Hamamelidaceae Hanguanaceae H

aptanthaceae Harmandaceae Hebenstretiaceae HectorellaceaeHederaceae Hedyosm

aceae Hedyotidaceae Hedysaraceae Heisteriaceae Heleniaceae Heliamphoraceae H

elianthaceaeHelianthemaceae Helichrysaceae Heliconiaceae Helicteraceae Heliotropi

aceae Helleboraceae Heloniadaceae HelosidaceaeHelwingiaceae Hemerocallidaceae

Hemimeridaceae Hemiphylacaceae Henriqueziaceae Henslowiaceae Hermanniaceae

Hernandiaceae Herniariaceae Herreriaceae Hesperocallidaceae Heterantheraceae He

teropyxidaceae HeterostylaceaeHibbertiaceae Hibiscaceae Hilleriaceae Himantandra

ceae Hippocastanaceae Hippocrateaceae HippomanaceaeHippophaeaceae Hippurida

ceae Hirtellaceae Holacanthaceae Homaliaceae Hopkinsiaceae Hoplestigmataceae H

ordeaceaeHornschuchiaceae Hortensiaceae Hortoniaceae Hostaceae Houstoniaceae

Huaceae Huerteaceae HugoniaceaeHumbertiaceae Humiriaceae Hyacinthaceae Hyd

atellaceae Hydnoraceae Hydrangeaceae Hydrastidaceae HydrillaceaeHydrocharitacea

e Hydrocotylaceae Hydrogetonaceae Hydroleaceae Hydropeltidaceae Hydrophylacac

eae HydrophyllaceaeHydrostachyaceae Hymenocardiaceae Hypecoaceae Hyperanth

eraceae Hypericaceae Hypopityaceae HypoxidaceaeHypseocharitaceae

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Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta

ceae ImperatoriaceaeInocarpaceae Inulaceae Ionidiaceae Iriarteaceae Iridaceae Irvi

ngiaceae Isnardiaceae Isophysidaceae IteaceaeStrasburgeriaceae Ixiaceae Ixioliriace

ae Ixonanthaceae

Jalapaceae Jamboliferaceae Japonoliriaceae Jasionaceae Jasminiaceae Johnsoniaceae

Joinvilleaceae JuglandaceaeJulianiaceae Juncaceae Juncaginaceae Juniperaceae Jus

siaeaceae Justiciaceae

Kadsuraceae Kaliphoraceae Kalmiaceae Kaniaceae Kiggelariaceae Kingdoniaceae Kin

giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae

Krameriaceae

Labiatae Lacandoniaceae Lachenaliaceae Lacistemataceae Lactoridaceae Lactucaceae

Lagerstroemiaceaee LamiaceaeLampsanaceae Lanariaceae Langsdorffiaceae Lantan

aceae Lapageriaceae Lardizabalaceae Lasiopetalaceae LauraceaeLawsoniaceae Laxm

anniaceae Lecythidaceae Ledaceae Ledocarpaceae Leeaceae Leguminosae Leitneriac

eae LemnaceaeLennoaceae Lentibulariaceae Lentiscaceae Leoniaceae Leonticaceae

Lepidobotryaceae LepidocarpaceaeLepidocaryaceae Lepidocerataceae Leptaulaceae

Leptospermaceae Lepturaceae Lepuropetalaceae LeuctenbergiaceaeLibocedraceae

Licaniaceae Ligustraceae Lilaeaceae Liliaceae Limeaceae Limnanthaceae Limnocharit

aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace

ae Linderniaceae Lindleyaceae Linnaeaceae LippayaceaeLiriaceae Liriodendraceae Li

ssocarpaceae Littorellaceae Loasaceae Lobeliaceaee Loganiaceae LomandraceaeLoni

ceraceae Lopeziaceae Lophiocarpaceae Lophiolaceae Lophiraceae Lophophytaceae L

ophopyxidaceae LoranthaceaeLotaceae Lowiaceae Lupulaceaee Luxemburgiaceae Lu

zuriagaceae Lychnidaceae Lyciaceae LyginiaceaeLygodisodeaceae Lysimachiaceae Ly

thraceae

Macarisiaceae Mackinlayaceae Madiaceae Maesaceae Magnoliaceae Malaceae Male

sherbiaceae MalortieaceaeMalpighiaceae Malvaceae Mangiaceae Manicariaceae Ma

paniaceae Marantaceae Marathraceae MarcgraviaceaeMartyniaceae Mastixiaceae

Matricariaceae Maundiaceae Mayacaceae Medeolaceae Medusagynaceae Peridiscac

eaeMelaleucaceae Melampyraceae Melanophyllaceae Melanthiaceae Melastomatac

eae Meliaceae Melianthaceae MelicaceaeMeliosmaceae Melittidaceae Melochiacea

e Memecylaceae Mendonciaceae Menispermaceae Menthaceae MenyanthaceaeMe

nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

47

Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

Nypaceae Nyssaceae

Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

ceae Papilionaceae Pappophoraceae PapyraceaeParacryphiaceae Parasitaxaceae Par

ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

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hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

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cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

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ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 3: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

3

from each family to the more detailed treatments on the individual order pages In

larger families I tend to focus on literature that deals with monophyletic groups that

include fifty or more taxa in smaller families the coverage is more detailed

I have emphasized plant families because they are the groups - admittedly partly

arbitrary as to circumscription but now for the most part monophyletic - around which

many of us organize our understanding of plant diversity I also pay attention to

groupings of families because so much progress has been made in the last decade in

particular in sorting them out Infrafamilial groups in groups like Poaceae Malvaceae

and Ericaceae are also included and these are being added to as studies become

available

But we dont want to know just about clades we want to know what makes clades

unique the synapomorphies or shared derived characters of those clades that first

appeared in their immediate ancestors However for the most part our knowledge of

synapomorphies remains poor ss we will see finding out the composition of clades is

often easier than finding the synapomorphies for the same clades (see the discussion

below) And of course knowing about synapomorphies is just one aspect of

understanding the whys and wherefores of the evolution and diversification of seed

plants our ultimate goal

ON CLASSIFICATIONS IN GENERAL AND THIS CLASSIFICATION IN PARTICULAR

On classifications in general

Classifications in the broad sense are box-in-box group-in-group or partwhole

naming devices that we use to communicate aspects of our knowledge of things in

general From this point of view there is nothing particularly special about biological

classifications apart perhaps from their sizes For any biological classification system to

be effective it must be stable universal (ie be used by a wide range of people)

comprehensive (if too many taxa are unplaced at the level of interest the classification

is of less utility) and it must enhance communication of knowledge by helping us to

relate things (Stevens 2006a for references) Phylogenetic classifications convey aspects

of our knowledge about phylogenetic relationships of organisms Thus a family is clearly

flagged as such and is a monophyletic group that can contain several genera also

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

4

flagged as such and also monophyletic but a genus can never include families Generic

family etc names are simply words we use to denote appropriate parts of phylogenies

and minimal aspects of their relationships and the irreducible aspect of relationships

that is emphasized here is monophyly A monophyletic group is one containing only and

all descendents of a common ancestor and it is often characterizable by apomorphies

that is shared derived characters that appeared in the ancestral lineage or stem clade

of that monophyletic group

Thus I am using a flagged ie ranked hierarchy for naming taxa rather than the

unranked systems that have recently been proposed (see below) The rank terminations

used (-ales -aceae etc - the flags) merely suggest relative positions of groups in the

local hierarchy If Ericaceae and Vaccinioideae are part of the same monophyletic group

the latter must refer to a clade nested within the former even if neither can necessarily

be directly compared with Polemoniaceae and Cobaeoideae (other than all being

putatively monophyletic groups) Such a flagged hierarchy is useful as a mnemonic and

communication device (eg Stevens 2006a) It improves memorization and emphasis on

families and orders as here is a didactic device - families are monophyletic units useful

in communication major units learned by biologists and others world-wide

The distinction between grouping and ranking is extremely important as is how we

interpret the latter We can both agree that there is a genus Acer yet disagree as to

whether it should be in Aceraceae or submerged in Sapindaceae Although from one

point of view this disagreement is utterly trivial it can have profound consequences if

we misunderstand the nature of the classificatory hierarchy Taxa at the same rank are

equivalent only by designation and have nothing necessarily in common (unless they

are sister taxa) other than their monophyly Rank as used here has no meaning other

than signifying a monophyletic group that includes other monophyletic groups with

appropriately subordinate rank terminations Taxa at the same rank have often been

treated incorrectly as if they were equivalent by biologists attempting to understand

evolutionary or biogeographic problems (see Bertrand et al 2006 for detailed

discussion) even if those constructing or using classifications - including Darwin (1859) -

have been explicit about the non-equivalence of taxa at the one rank (Stevens 1997) In

fact rank terminations have relatively infrequently been used by taxonomists to reflect

absolute rank although Linnaeus (at least in theory) at the level of genus and species

may be such an example (Classifications where rank is absolute taxa at the same rank

somehow being comparable entities are class hierarchies in the strict sense - Stevens

2002 2006a) It has also been suggested that taxon rank be adjusted so that rank

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

5

somehow reflects the degree of morphological differences between taxa or that taxa at

the same rank be based on similar characters or show a similar amount of distinctness

This might be possible using phenetic methods of analysis but is very difficult if ones

classification is phylogeny-based as here it could promote instability if used in taxa

where such a criterion had not previously been used and it might also inadvertently

suggest that taxa might be equivalent (for an example see Fritsch et al 2008) There

have also been proposals that rank could reflect the age of the clade (eg Hennig 1966)

with clades that have diverged by a particular time all being given the same rank Apart

from the fact that aging times of divergence of clades is still a difficult enterprise huge

disruptions to our nomenclature would result Recent suggestions which invoke the use

of age in classifications focus on providing a standardized timeclip ie a set of letters

referring to a particular geological period that could simply be added to a conventional

taxon name (Avise amp Mitchell 2007) However even such timeclips are unlikely to come

into general use soon

It would be impossible even to think about a higher-level classification such as this

without the advances in our understanding of relationships made by the phylogenetic

analyses of molecular data carried out over the last twenty years One can then

integrate the data to be found in both classical and recent morphological studies with

these phylogenies For the dramatic changes in this area see for instance the

pessimistic attitude towards orders in Davis and Heywood (1963 107-108) The most

unsatisfactory taxon in Angiosperm classification they were indefinable their

circumscription was not fixed etc Families they thought were likely to be the largest

natural unit within the mono- or dicotyledons Along the same lines almost three

quarters of the orders (4459 monofamilial orders ignored) recognised by Cronquist

(1981) are not monophyletic ie they do not contain all and only the descendents of a

common ancestor Most of those orders that are monophyletic are very small

(Zingiberales with eight families are the largest) for families on the other hand about

two thirds (189273) are monophyletic

Turning now to phylogenetic classifications and the particular classification used

here Backlund and Bremer (1998) provide a useful discussion on the principles of

phylogenetic classification that is applicable at all levels apart from species (see also

Stevens 1998 also Albach et al 2004 Entwisle amp Weston 2005 Pfeil amp Crisp 2005 etc

for examples) Backlund and Bremers main principle is that taxa that are recognised

formally should be monophyletic However this does not indicate which particular

clades we might wish to name as families genera etc and talk about in general

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

6

conversation If a well-supported hypothesis of monophyly is a necessary prerequisite

for a group to be named it is not a sufficient prerequisite (but cf the PhyloCode -

Cantino amp de Queiroz 2006 Cantino et al 2007) Not all clades need be named indeed

it would barely be practicable (or practical) to do this To decide which clades should be

named additional criteria can be invoked Other things being equal it is helpful if 1 taxa

formally recognised are easily recognizable 2 groups that are well-established in the

literature are preserved 3 the size of groups is taken into account and 4

nomenclatural changes are minimized (Backlund amp Bremer 1998) Thus numerous small

groups have little to recommend them since individually they summarize little

information and tend to clog the memory while groups that are too big may be

amorphous Somewhat similarly Godfray and Knapp (2004 p 562) note that users

want stable informative and accessible classifications that enable easy identification

(see also Simon 2008) - although invoking users without specifying those who make up

this group is not very helpful This classification is for all interested in comparative

biology hence the emphasis on monophyly Although the clades named are sometimes

difficult to characterise there are as we shall see many ways of making such a

classification accessible to all

Problems with this emphasis on monophyly may be caused by reticulation events

such as hybridization endosymbiosis and lateral gene transfer but they are unlikely to

be common confusing factors here Genera can often be pegged to above the level at

which hybridization is at all common However in Poaceae-Pooideae-Triticeae there are

some intractable problems where extremely well established common usage and the

principle of monophyly are likely to remain at odds Many genera are certainly not

monophyletic here being allopolyploids and the genera are ultimately based on the

different genomes they contain (Dewey 1984 Loumlve 1984 Barkworth 2000 for a history

of Triticeae classification Petersen et al 2006) There is also extensive reticulation

reported within Danthonioideae (Pirie et al 2009) Evidence also increases of old

hybridization events elsewhere in flowering plants that at the very least cause

discordance between relationships suggested by different genomic compartments as in

Smedmark and Anderberg (2007 Sapotaceae) and Fehrer et al (2007) Morgan et al

(2009) and Pelser et al (2008 all Asteraceae - hybridization is likely to be quite a

problem at the generic level here) and genera like Medicagoare turning out to have

highly reticulating relationships at the species level (Maureira-Butler et al 2008) Of

course there are many problems trying to apply the concept of monophyly to species

and for many - but not all - biologists strict monophyly is a less important criterion at

this level (Funk amp Omland 2003 see much of the discussion in Houmlrandl 2006)

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

7

The major endosymbiotic events that characterize the clade of which flowering

plants are a part (and gave rise to chloroplasts and mitochondria) are very ancient and

cause no problems for the student of multicellular organisms However lateral gene

transfer has been detected in a number of situations between quite unrelated

organisms (eg Bergthorsson et al 2003 - Amborella and liverworts see also below but

cf Goremykin et al 2009) and it may be particularly common in mitochondria (Sanchez-

Puerta et al 2008) Here too there are no major problems providing one is careful

such transfers do however raise all sorts of interesting biological questions (see

Richardson amp Palmer 2007 for a summary) Although there is increasing evidence for the

importance of genome duplications - hybridization is one cause of this - at various times

during the evolution of seed plants and of palaeopolyploidy events within eg the

Lauraceae and Magnoliaceae clades (Soltis et al 2009 for a summary see eg

the Characters page for further discussion) these too do not currently seem to pose

problems for the adoption of monophyly as the sine qua non of groups to be recognised

formally in this phylogenetic classification but it can make detecting orthologous genes

difficult

The accessory principles of Backlund and Bremer (1998) should be used in

combination Thus keeping the monogenericPlatanaceae separate from its sister

taxon Proteaceae is justifiable Both are much-used names that signal well supported

well defined and easily recognisable groups that have long been recognised as distinct

have several synapomorphies and do indeed look very unlike each other Combining

the two would yield a clade with few obvious apomorphies not to mention the fact

that Nelumbonaceae should by the same logic (it is also monogeneric) also be included

in the expanded family On the other hand it is difficult to justify the continued

recognition of Callitrichaceae or Hippuridaceae monophyletic and distinctive although

they may be If they were recognised several poorly characterised clades would also

have to be carved out ofPlantaginaceae in any classification that aimed to convey a

comprehensive view of the worlds flora The continued recognition

of Valerianaceae and Dipsacaceae also tends to run into this problem (see also Pfeil amp

Crisp 2005 Orthia et al 2005 Albach 2008 etc for useful practical discussions of such

matters) But there are no absolute guidelines If Podostemaceae turn out to be sister

to Hypericaceae (for references here and elsewhere in the Introduction see the

individual families) the subsequent moderate dismemberment of Clusiaceae sl is not

be too high a price to pay for the continued recognition of Podostemaceae Hence the

somewhat provisional recognition of Hypericaceae and Calophyllaceae as well as

Clusiaceae below the families can all be recognized and the name Podostemaceae in

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

8

particular is very well established In a somewhat similar situation if Lemna and its

relatives are a clade sister to most other Araceae should they be recognised as a

separate family Gymnostachys a phenetically fairly distinctive taxon as well as the less

phenetically distinct Orontioideae would have to be recognised as a separate families

(or combined as a single family) too but Araceae in a somewhat restricted sense would

be somewhat more morphologically coherent although not greatly so and not notably

distinct However it is in the very nature of such decisions to be somewhat arbitrary

and unsatisfactory hence the emphasis on consensus classifications here and on

classifications as simply being a means to an end

A useful distinction can be drawn between crown groups and stem groups The

former are monophyletic and include the extant members of a clade and their

immediate common ancestor (see the figure below) The groups characterized in this

site are such groups Thus Proteaceae here are crown group Proteaceae apomorphies

like the single carpel four-merous perianth etc being found in this common ancestor

Stem groups on the other hand include all the members of a lineage immediately after

its split from its sister group and all branches of this lineage In the case of Proteaceae

the stem group would include everything after its split from its sister group

Platanaceae Obviously most of the organisms in the lineage that terminates in crown-

group Proteaceae are unknown only a few being known as fossils and it is not known

exactly where particular apomorphies of crown group Proteaceae evolved along this

lineage

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

9

Indeed there is no reason other than convention or convenience why any group

should not be segregated into several smaller monophyletic groups or merged to

produce a larger unit we can talk about one large thing or about several smaller things

Thinking about aspects of size findings in ethnobiology and cognitive psychology can be

used to suggest that a moderate number - probably fewer than 500 - of families is a

reasonable goal at which to aim and that groupings of taxa throughout any system

should be rather small in size (eg Berlin 1992 Stevens 1994 1997) Major systems such

as those of Linnaeus and Bentham and Hooker were constructed explicitly so as to ease

the burden on the memory (Stevens 1997 2002 see also Scharf 2007) the latter in

particular ensuring that all groups in their classification were relatively small often

containing three to eight immediately subordinate taxa - but by no means all their

groups were formally named Along the same lines Burtt (1977b) suggested that the

number of names at any rank should be at most one third those at the immediately

lower rank - and monotypic taxa might not need a formal name Consistent with such

ideas a fairly broad view of families and orders is taken here whenever the constraints

of monophyly and other criteria used when constructing classifications (see above)

permit The APG system is thus a convention consistent with ideas such as those

expressed by Backlund and Bremer (1998) for similar conventions see eg the Grass

Phylogeny Working Group (2001) and for fungi Hibbett et al (2007)

There are other views Thus Takhtajan (1997) has suggested that smaller families

are more natural This is incorrect Monophyletic groups that include fewer taxa -

Takhtajans smaller families - do not necessarily have more apomorphies than larger

groups even if members of smaller groups will always have more features in general in

common than the larger group that includes them That is they will have their

apomorphies their unique features as well as progressively more plesiomorphies

features found both in the small groups and in the larger clades of which they are a part

having more features in common is indeed one common meaning of more natural

(Note that the implication of the word natural has long been a group of the kind

[usually unspecified] that I think should be recognised and if other botanists had

different ideas too bad their groups were necessarily not natural [see Bather 1927] It

is thus rarely a helpful word and is not used here) Furthermore if this approach is

adopted we will find a slippery slope ahead By this kind of argument all families should

be very small since their members will have a great deal in common and so will be

most natural However as families (for example) are split the relationships that are

evident between the segregates and that were responsible for their being placed in a

single family in the first place will seem to necessitate the recognition of a new order

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

10

etc as is evident in Takhtajans own work - general taxonomic inflation is the result (see

also comparable suggestions in a cladistic context for Brassicales in particular - Ronse de

Craene amp Haston 2006) Such splitting is also questionable when teaching and learning

families since the student needs to understand the system as a whole However for

some genera removed from the families that until now have included them the

phenetic-classificatory-phylogenetic structure in their new home may mandate the

recognition of small families On the other hand Takhtajans suggestion that narrowly

defined families are more useful for phylogenetic studies may be true Indeed I have

more than once regretted prematurely combining groups whether species (in the

context of monographic work) or families (in the course of preparing these notes) but

this is largely a separate issue

Van Steenis (1978) Philipson (1987b) and others have rightly questioned the value

of splitting a group when ideas of the relationships of its constituent members have not

changed - that is very good reasons have to be provided for splitting a family if the

genera within it remain part of the same clade rather than belonging to another clade

Thus APG (2003) broadened the circumscription of Malvaceae because of the

parapolyphyly of some of the families that had historically been associated with it (Judd

amp Manchester 1997 Alverson et al 1999 Bayer et al 1999) These families particularly

Tiliaceae and Sterculiaceae were not at all easy to distinguish their close relationship

(see eg Brown 1814) and overall similarity had long been conceded and to some

workers at least their combination has come as something of a relief Although most of

the larger clades within Malvaceae sl remain difficult to distinguish even with flowers

Cheek (2007) opts for a wholesale and novel dismemberment into ten families

however the very good reasons for doing this are wanting

The same principles are of course applicable when it comes to dividing genera little

other than a headache is gained by splitting genera such

as Drosera and Gnetum (Doweld 2000) as has recently been proposed Thus if an

established genus divides into two (or more) clades this is not a signal for recognising

two groups at the same level - so here it could be argued that the dismemberment

of Pterostylis (Jones amp Clements 2002b) was somewhat unfortunate Along the same

lines if a newly-discovered taxon is sister to an existing named taxon - say a genus - this

does not necessitate the description of a separate genus for the newly described species

(cf Davis 2002) (Of course some [eg Thorne 1976] have suggested that the sizes of

gaps between groups at the same rank should be similar but any principle like this is

inherently flawed since morphological gaps are more unstable than phylogenetic

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

11

relationships and applying it across all flowering plants would both be difficult and cause

substantial changes - see also below) Humphreys and Linder (2009) provide a well-

documented survey of generic concepts in plants which the reader should consult they

note that generic limits (broad versus narrow) have oscillated historically and that

currently larger genera tend to be recognised because studies tend to be on a broader

scale than in the past

Note that invoking similarity or difference - whether qualified (considerable

similarities substantial differences) or not - in a cladistic context as justification for

combining or splitting taxa is not a particularly strong argument (see eg

CardiopteridaceaeStemonuraceae - Karingrehed 2002c) Similarity and difference can

neither be defined precisely since what may seem to be substantial similarities to me

may not to the next person nor are they likely to be stable in the face of our changing

knowledge of morphology and what might be synapomorphies

I might have prefered to merge some families recognised here or split others but

by and large I do not think my own preferences matter very much - and I take the same

position with regards to comparable preferences expressed by others Indeed the

bottom line is that in flagged hierarchies of the kind used here the limits of any

monophyletic unit generally taught and discussed particularly other than species can

be established only by convention and consensus (eg Stevens 2002 2006a Entwisle amp

Weston 2005) This is rather different from the reasonable observation that the

phylogeny itself cannot be achieved by consensus (Thorne 1976) for in most cases there

is a fact of the matter when it comes to relationships Given the increasing support for

the outlines of angiosperm phylogeny a stable consensus classification based on this

phylogeny seems attainable Indeed in addition to providing current ideas of

relationships of seed plants in a synthesised form this site is part of an attempt to build

such a consensus about the circumscription of taxa (see APG 1999 2003 2009 Grass

Phylogeny Working Group 2001 Mabberley 2008) Reaching such a consensus is vital

since what we know of angiosperm phylogeny allows a very large number of

classifications to be based on it and as we find out more the number of possible

classifications increases greatly Unfortunately however nature does not dictate what

the classification should be All classifications are constructed by humans to

communicate particular aspects of groups and relationships Our goals as systematists

are surely to produce robust hypotheses of relationships to understand the evolution of

morphology and the like - but not to argue ad nauseam whether something should be

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

12

a family or a subfamily That way surely lies madness and worse the discredit of our

discipline

There are similar issues whatever naming system is used Thus in phylogenetic

naming (Baum et al 1998 for an example but cf Baum et al 2004 for the PhyloCode

see Cantino amp De Queiroz 2006) an unflagged hierarchy is used in which any

terminations of names used are uninformative about the relative position of taxa If one

adopts the principle of phylogenetic naming one indeed does not have to worry about

the nomenclatural consequences caused by lumping or splitting any well-supported

clade can be named without affecting the name of more or less inclusive clades

Unfortunately unflagged hierarchies have very serious deficiences as communication

devices because they lack one aspect essential in language biological or otherwise -

they contain no intrinsic information about the relationships of the group in question to

others (eg Pfeil amp Crisp 2005 Stevens 2006a) Recent suggestions for using prefixes like

Apo- and Pan- to PhyloCode names will however allow limited information of this

kind to be conveyed but only as it pertains to individual branches and current

proposals do not even mandate that the prefixes be employed consistently In any

event such proposals simply prevent the potential tripling of the number of quite

different names used to describe different aspects of a phylogenetic tree over those

used to name monophyletic groups pure and simple In general where n is the number

of extant species in a group the number of clades in such a group = n-1 (Species will

also need names too for their names see Dayrat et al 2008) Importantly here too

consensus over the clade names commonly learned by students and used in herbaria is

needed otherwise communication will be impeded the names themselves will provide

no guidelines as to which should be chosen The situation is of course more complicated

than this Terminations that convey ideas of rank in a phylogenetic classification can also

be used in phylocode names - however there they will carry no implications of rank

How they will be used is another matter of course

Of course there are other philosophies of classification and some still prefer

evolutionary classifications There classificatory principles differ substantially from those

followed here eg the recognition of paraphyletic taxa may be permitted however

detailed reasons for prefering the taxa that are recognised are rarely given although

nature and natural groups are often mentioned (cf Stuessy amp Koumlnig 2008) For

summaries of commonly used systems see Brummitt (1992) and Mabberley (2008)

new evolutionary systems appear every year or so However even those who allow or

promote the recognition of paraphyletic groups (eg Grant 2003 Thorne 2007

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

13

Heywood et al 2007) may find it of interest to examine a system recognizing only

monophyletic groups it provides a rather different understanding of evolution

Evolutionary classifications in general try and combine phylogeny and

morphological gaps although that is no easy thing to do - it is akin to combining chalk

and cheese (for an attempt to make this impossible task seem to be more objective see

Stuessy amp Koumlnig 2008) However there is a principle from evolutionary classification

that is relevant and I think quite useful here The size of the gap between two groups

tends to be inversely proportional to the sizes of the groups involved (Davis amp Heywood

1963) One can imagine a situation in which a large group is formally divided even

although the distinguishing characters of the two are weak whereas a smaller group

similarly divisable will be left intact

To summarize If hypotheses of phylogeny remain stable we should be able to base

a stable classification on that phylogeny and then get on with our work that is testing

the phylogenies we have elucidating phylogenies in areas where relationships are

unclear studying the evolution of morphology describing species etc In this context

the spread of the Angiosperm Phylogeny Group system (see below) and its widespread

utilisation in technical literature also floras (eg van der Meijden 2005) textbooks (eg

Simpson 2006 Judd et al 2007 [third edition]) dictionaries (Mabberley 2008) more

popular literature (eg Souza amp Lorenzi 2005 Spears 2006) and as an outline for a new

herbarium sequence (Haston et al 2007) is gratifying The posibility that one might be

able to develop a stable phylogeny-based classification of families and in particular

orders represents a dramatic turn-around from the pessimistic attitude about such

higher-level groupings expressed by Davis and Heywood (1963) and Thorne (1976) the

latter even suggesting that we should bury forever the metaphor of the phylogenetic

tree as highly unrealistic (ibid p 56) Returning to Godfray and Knapps (2004) users

of classifications who want a stable informative and accessible classification that

enables easy identification - unfortunately they want cake with everything and cannot

get it - these pages attempt to satisfy as many of their needs as possible but phylogeny

and monophyly are the primary shapers of the APG classification

On this classification in particular

Here I very largely follow the Angiosperm Phylogeny Group classification (APG

2003) Any differences are not to be interpreted as differences in principle simply that

new phylogenies continue to be published and here I attempt to provide an overview

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

14

of current ideas of higher-level relationships of seed plants The Angiosperm Phylogeny

Group classification is based on well-supported relationships evident in the numerous

molecular studies that began to appear in the late 1980s (see APG 1999 for the

principles underlying the classification) but recent phylogenetic work does not

contradict the major outlines of the trees used by APG II (2003) or even those in APG

I (APG 1999) The APG classification has for the most part been conservative and I

am similarly conservative here For the names and authorities I follow APG III (2009)

although the names of the authors may not always be correct these being in something

of a state of flux for the superordinal names used see Chase and Reveal (2009)

although classification at this level here is not exhaustive - only clades with several

orders have superordinal names

Some changes in our ideas of relationships and hence in the clades we talk about

are particularly likely in parts of Lamiales Caryophyllales and Malpighiales for example

although this will not affect the orders themselves Thus some changes to clade

circumscriptions are to be expected although I do not expect them to be substantial

but changes are neither a defect of cladistics nor a necessary consequence of the use of

molecular data Clades are hypotheses of relationships and as hypotheses they may be

overturned However work since APG I has not suggested other than minor changes

in the compositions of the orders even if the odd genus or even family is turning out to

be seriously misplaced - recent examples are Hydatellaceae (from monocots-Poales to

Nymphaeales Saarela et al 2007) Guamatelaceae (from Rosales-Rosaceae to

Crossosomatales Oh amp Potter 2006) and Perrottetia and Bhesa (from Celastrales-

Celastraceae to Huerteales and Malpighiales respectively Zhang amp Simmons 2006) The

main changes have been clarification of the relationhips of individual families or groups

of families that were of uncertain position eg of Chloranthaceae (Moore et al 2007)

Ceratophyllaceae (Jansen et al 2007) and unplaced asterid II families (Winkworth et al

2008a) As mentioned above phylogenies in the area of main interest in these pages are

overwhelmingly tree-like Although there is evidence for acquisition of host plant genes

by parasites (see below) even wider but inexplicable transfer of mitochochondrial

genes in plants like Amborella (Bergthorsson et al 2004 cf Goremykin et al 2009a)

and even transfers of nuclear genes (Vallenback et al 2008) these are the exception

rather than the rule

In cases where the Angiosperm Phylogeny Group II allowed alternatives as to the

limits of families - Papaveraceae in the broad sense or Papaveraceae plus

Pteridophyllaceae plus Fumariaceae Proteaceae in the broad sense or Proteaceae plus

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

15

Platanaceae - the choices made here follow common usage eg as in textbooks like

Judd et al (2007) and Simpson (2006) and particularly in the new edition of

Mabberleys The Plant Book (Mabberley 2008) This latter is itself an attempt to reflect a

consensus the result of taking the opinions of botanists at several meetings A largely

similar consensus is reflected in the APG III classification (APG 2009) where you will

see that alternative classifications have been dispensed with For many people the

existence of such alternative classifications will simply confuse so agreement over

which groupings to use when alternatives are permitted and then dispensing with the

alternatives that are not commonly used seems reasonable I am making the few

changes that are needed to bring the classification here in line with these others but

only gradually since there is not enough time in each day

As already mentioned higher-level relationships in general and the composition of

orders in particular have always presented something of a challenge to systematists

(eg Davis amp Heywood 1963) That being said the composition of some clades like

Apiales Crossosomatales and Pandanales is decidedly unexpected However it is

interesting to see that these higher level clades are generally accepted even in works

with different classificatory philosophies (eg Heywood et al 2007) For clades like

Malpighiales many of the family groupings within Asparagales etc attempts to find

distinctive characters have largely failed (but see Endress amp Matthews 2006a also

the Apomorphies page here) Interestingly as with families some groupings suggested

by molecular studies are supported by morphological andor chemical characters that

have long been known sometimes for over a hundred years the relationship between

Pittosporaceae and ApiaceaeAraliaceae are a case in point (Hegnauer 1969b and

references) As our knowledge of morphology and chemistry improves we can hope for

improvements in the characterisations of clades at all levels

In many cases the new family limits of the Angiosperm Phylogeny Group (see

APG 1999 2003) are not really controversial although changes from the limits

commonly accepted only a decade ago are sometimes quite dramatic (eg Wagenitz

1997) Thus the split of the old Saxifragaceae sl is necessitated by its extreme

polyphyly as also with Icacinaceaesl and Cornaceae sl However the limits and

relationships of such groups had long been considered to be unsatisfactory but until

recently there had been no real reason to prefer one proposed arrangement over

another It is generally accepted that the limits of Lamiaceae and Verbenaceae have to

be redrawn although there is as yet no compelling evidence that the redrawn taxa are

not sister taxa (there is no evidence that they are) But whatever their relationships the

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

16

content of the clades has changed considerably and incidentally they are now easier to

identify than before the decision to recognise two families is not difficult The same is

true for Salicaceae and Achariaceae (Malpighiales) two previously small families that

have received the bulk of the old Flacourtiaceae Clade and hence taxon limits remain

difficult around Theaceae (Ericalessee alsoSladeniaceae Pentaphylacaceae)

and Euphorbiaceae (Malpighiales see

also Phyllanthaceae Picrodendraceae Putranjivaceae etc) Here current groupings

may not represent quite such dramatic changes in our understanding of relationships

Indeed parts of the old Euphorbiaceae that were separated may yet go back together

particularly Phyllanthaceae and Picrodendraceae (see Wurdack et al 2004 Davis et al

2005) even if Theaceae should indeed be dismembered (Geuten et al 2004) However

given that there is no molecular evidence that currently warrants combining all the

segregates of Euphorbiaceae and even if some do come together the clades the

families represent suggest novel groupings not recognised in current classifications

Furthermore Rafflesiaceae appear to be embedded within Euphorbiaceae s str (Davis

et al 2007) so the family is divided Of course maintaining even a moderately narrowly-

circumscribed Euphorbiaceae would entail reducing the iconic Rafflesiaceae to

synonymy Relationships in core Caryophyllales especially

around Phytolaccaceae andNyctaginaceae and also Portulacaceae are incompletely

understood and refashioning of taxon limits will doubtless be needed as cladistic

relationships become apparent (eg see Nyffeler 2007 Ogburn 2007a b) Some

groupings in the old Icacinaceaeand Olacaceae also remain unclear

The discovery of the relationships of parasitic and aquatic groups have presented a

particular challenge to systematists Morphologically some of these plants are so highly

modified that interpretation of the plant body in conventional terms is difficult or even

impossible Thus parasitic groups (broad-leaved angiosperms only) such as Rafflesiaceae

are hard to place since both the vegetative body and the flowers are changed almost

beyond recognition (flowers of rafflesiaceae are a good example of gigantism - Davis et

al 2007) Furthermore plastid gene sequences may be difficult or impossible to obtain

the chloroplast DNA in particular being highly degraded and the rate of molecular

change in general may be high so the problem of long-branch attraction is serious (eg

Duff amp Nickrent 1997 Nickrent et al 1998 Caddick et al 2002a G Petersen et al

2006b) Echlorophyllous sapromycoheterotrophic taxa mostly known from monocots

present similar problems However progress is being made placements for

Apodanthaceae Rafflesiaceae Mitrastemonaceae Cytinaceae and Cynomoriaceae have

recently been suggested (Barkman et al 2004 2007 Davis amp Wurdack 2004 Nickrent et

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

17

al 2004 Davis et al 2007) it seems likely that Burmanniaceae sl are polyphyletic but

both parts are in Dioscoreales (Merckx et al 2006) and relationships within the largely

hemiparasitic Santalales are also gradually being clarified (Maleacutecot 2002) The inclusion

of parasitic taxa in general molecular analyses can cause conniptions (eg Nickrent et al

2004 Davis et al 2004 Chase et al 2006 G Petersen et al 2006b) and there can be

horizontal transmission of genes (eg Davis amp Wurdack 2005 Vitaceae to Rafflesiaceae

Barkman et al 2007 the mitochondrial atp1 gene commonly moves)

Similarly in water plants neither vessels in particular nor much xylem in general is

needed leaves are highly modified and water-mediated pollination if adopted may

well be associated with major changes in floral morphology Here too recent molecular

studies suggest that aquatic groups with hitherto problematic relationships may find

homes Thus Podostemaceae are close to Clusiaceae and Calophyllaceae and are sister

to Hypericaceae (Malpighiales Kita amp Kato 2001) Hydatellaceae which used to be in

Poales are part of Nymphaeales (Saarela et al 2007) and Hydrostachyaceae may be

close to Hydrangeaceae (Cornales Xiang et al 2002) or perhaps should be placed in

Lamiales note that in the first two cases in particular there are morphological and

chemical features that support such a move If Podostemaceae are indeed close to

Clusiaceae I look forward to seeing hypotheses to explain how the dramatic changes in

the vegetative body that have made Podostemaceae so problematic for generations of

systematists took place That conventional wisdom has trouble in understanding or

explaining how the morphologies of groups like Clusiaceae and Podostemaceae can be

related is largely a problem with conventional wisdom Ceratophyllaceae seem to be

finding a position as sister to eudicots (see Moore et al 2007) but their morphology is

so derived that there is no morphological evidence of which I am aware for this

relationship There are also similar although less extreme morphological problems with

many plants pollinated by wind

In such situations especially with some water plants mycoheterotrophs and

parasites a variety of characters may be affected and particularly under such

circumstances groups have tended to be formed using a few or even only a single

character that seems to provide evidence of relationships Add to this the tendency to

weight some characters particularly strongly on a priori grounds and the result was

either the recognition of conglomerate taxa such as Amentiferae which now appear to

be highly polyphyletic or the segregation of families like Plantaginaceae s str (now

much expanded) and Leitneriaceae (now in Simaroubaceae) - all of these are more or

less wind pollinated

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

18

Even although relationships of many parasitic and aquatic groups are being

clarified and at least some of the proposed phylogenetic positions seem to be well

supported it is indeed often still very difficult to understand how such groups have

evolved from their more morphologically conventional relatives However we have had

all sorts of preconceptions as to how characters should evolve Thus we used to

assume that features like highly scalariform vessel perforation plates or the complete

absence of vessels or a flower with an androecium that had many stamens a superior

ovary or separate petals were necessarily plesiomorphic or primitive and conversely

a simple perforation plate and the presence of vessels an androecium with few

stamens inferior ovary or petals that were connnate were almost necessarily

apomorphic or advanced (but cf eg Stebbins 1951) Such assumptions are incorrect

(eg Soltis et al 2005b) Carpels may become secondarily free carpels may fail to close

the seeds then developing outside the confines of the carpel as in some Aspagaraceae-

Nolinoideae Violaceae Berberidaceae Malvaceae-Sterculioideae etc) in Peliosanthes

teta perhaps the only species in Peliosanthes(Asparagaceae-Nolinoideae) the ovary

varies from superior to inferior (Jessop 1976 species limits here need close

investigation see also Kuzoff et al 2001 and Soltis amp Hufford 2002 Saxifragaceae

Apiales etc) many-seeded carpels can evolve from few-seeded carpels

(Razafimandimbison et al 2008) monoecy may be derived from dioecy (Schaefer amp

Renner 2010 and references) Classic studies such as those by Babcock (eg 1947)

on Crepis that assumed that evolution - in this case of the karyotype in particular - was

unidirectional have needed comprehensive re-evaluation (Enke amp Gemeinholzer 2008)

Most if not all characters have reversed andor evolved in parallel as is clear in the

discussion of ovary position here especially in Poales Saxifragales and Asparagales and

also in comments on such morphologically distinctive taxa (when compared with their

immediate relatives) such as Menyanthaceae and especially Pittosporaceae Parallel

evolution may occur even at the level of amino acid substitution as in the independent

acquisition of the phosphoenolpyruvate carboxylase (pepC) gene in C4 photosynthesis in

grasses (Christin et al 2007b see also Blaumlsing et al 2000)

Thinking of how such characters have been used in the past in classifications at the

generic level the results have been similar to those just discussed above for parasitic

plants and others Relying too much on animal pollination syndromes to mark generic

boundaries has all too often led to taxa that are highly unsatisfactory phylogenetically

(see eg Acanthaceae Bignoniaceae Campanulaceae Ericaceae Lamiaceae

Orchidaceae) and over-reliance on characters of fruit and seed (see eg Brassicaceae

and Apiaceae) has also led to unsatisfactory generic limits Again the more general

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

19

problem is the use of one or a very few characters that have been weighted a priori to

structure classifications (see also Garciacutea et al 2009) although it is perhaps not

surprising that such an approach sometimes results in the delineation of taxa that have

indeed turned out to be quite distinct (cf eg Burtt Astragalus versus Oxytropis)

Taxa recognized here are monophyletic yet many are polythetic at the

morphological level that is they lack unique features characterizing (defining) all and

only members of that family they can be recognised phenetically only by the unique

combinations of characters that they posess This is the result of evolution any

synapomorphy characterizing a taxon may be lost or modified beyond easy recognition

in some of its members or the synapomorphy may appear to be identical to a feature

that has evolved in parallel in a quite unrelated plant That plant groups are polythetic is

almost as much a feature of monophyletic taxa as the paraphyletic taxa common in

evolutionary classifications Some families now include substantial variation as

phenetically distinct derived groups are placed in their proper phylogenetic position -

examples are Ericaceae which include the erstwhile Empetraceae a wind-pollinated

group that is florally very different and the overwhelmingly large-flowered and animal-

pollinated Plantaginaceae which now include several derived small-flowered aquatic

and wind-pollinated groups that were previously placed in separate families

Thus some families as delimited here may not be easy to recognize However

remember that detecting relationships - use whatever characters you can even if they

are not obvious - and naming a plant - focus on easy-to-see characters that may not

reflect relationships - are quite different problems Taxa although natural may not be

readily recognizable indeed it was in exactly this context that Lamarck worked out the

basic principles of writing dichotomous keys in 1778 (see Scharf 2007 for keys and the

like) Of course Lamarcks idea of nature was very different from ours - he thought that

there was some kind of continuum of form on which living organisms were to be

situated with no real gaps anywhere - but this meant that his genera (for example)

might well not be sharply distinct from each other even if each were part of the real

continuum that was life Lamarcks separation of the establishment of relationships and

groupings in a system and identification of the plants in that system have been widely

accepted (but cf eg Godfray amp Knapp 2004 many of the contributors to Heywood et

al 2007) Perhaps the best way of identifying plants at the family level is by well-made

multiple access keys as in Watson and Dallwitz (1992a onwards family limits there may

differ substantially from those adopted here) Multiple access keys free users from the

constraints of dichotomous keys in which particular characters are needed at each step

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

20

of the identification process before there can be further progress Instead those

characters that happen to be evident on a specimen can be used in whatever order is

convenient when linked to illustrations glossaries etc their power is enormous (see

Dallwitz et al 2000 [2006] for the principles underlying their construction and use)

Nevertheless dichotomous keys such as those of Hutchinson (1973) and Franz Thonner

(Geesink et al 1981) have their uses Of course taxa coming out adjacent in keys may

well not be at all related

(When identifying large numbers of plants even more efficient than either style of

identification and certainly lots more fun is sight identification Unless you have a

photographic memory you have to build up your knowledge of comparative plant

morphology - on which the ability to make accurate identifications depends - by

repeated observation When faced with an unknown plant I always look for leaf teeth

and stipules distinctive hair types especially stellate and T-shaped hairs glands and

punctations of any sort the presence of latex or other exudate and check leaf insertion

smelling crushed leaves can also be helpful In this context nodal anatomy can usually

be checked using a razor and a hand lens or even simply carefully examining leaf scars

The short paragraphs added after most families may help in confirming familial

identifications)

For the record and for the little that it is worth there are 4 orders and 13 families

of gymnosperms characterised on these pages and together they include some 82

genera and 947 species For angiosperms comparable figures are 56 orders 445

families 13208 genera and 261750 species (of which monocots include 11 orders 89

families 2759 genera and 52760 species) Note however that higher mathematics

was never my strong point and anyway these are pretty meaningless figures even for

species which many (but not all) might concede smacked slightly more of reality than

other taxa estimates range as high as 422000 (Govaerts 2001) Furthermore numbers

of genera and species change daily Nevertheless as emphasized here families are

useful in teaching we as a community can ensure that their limits remain largely stable

and by concentrating on relatively few of them one can gain some familiarity with much

of the worlds flora For summaries of the sizes of orders and families and of the general

arrangement followed in these pages see also the Statistics page itself

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

21

A PHYLOGENETIC TREE OF SEED PLANTS (THE MAIN TREE)

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

22

The Main Tree above is a conservative summary of well supported relationships

between the major clades of extant seed plants Reading - interpreting and

understanding - such a phylogenetic tree is not necessarily easy so reading a tree may

require a bit of practice OHara (eg 1988 1992) provides an accessible introduction to

thinking and talking about history in the context of looking at phylogenetic trees More

recently Krell and Cranston (2004) Crisp and Cook (2005) and others have emphasized

how careful one must be when interpreting and talking about evolution in ladderized

trees in particular and phylogenetic trees in general Gregory (2008) provides a

particularly detailed discussion of this problem In ladderized trees the smaller (in terms

of numbers of terminals) sister taxon is consistently shown on the same side at every

node thus the trees tend to be pectinate like the teeth of a comb however the

horizontal axis that is as a result so evident has no polarity or particular direction The

use of the adjective basal when discussing phylogenetic trees is especially dangerous

(see also Wojciechowski et al 2004 D Soltis et al 2005b) When I use the term and the

context is not otherwise clear I am referring to the pectinations at the base of a

ladderised tree Remember that when talking about sister taxa one can never be basal

to the other or older or younger although members of one can be more derived - in the

sense of having more apomorphies - than the other (but even then one has to be

careful) The words primitive and advanced and lower and higher should

practically never be used especially when talking about taxa Amborellaceae and

Pinaceae are sister to all other flowering plants and Pinales and are basal to the crown

groups of all other flowering plantsPinales respectively but that does not mean they

are thereby primitive Pinaceae in particular have numerous apomorphies The word

plesiomorphic is far less loaded than primitive and can be used to talk about

individual characters

Most trees here have been more or less ladderized as is the one above The

ladderization may be imperfect for example you can see that on the Main Tree

asterids with ten orders follow rosids and relatives which have seventeen although

this is in part because of the currently unresolved nature of relationships between the

main clades of core eudicots (this is likely to change soon) The Main Tree could have

been drawn with Amborella Acorus or a host of other taxa at the far right without

offending any relationships phylogenetic trees are like mobiles the only fixed points

being the nodes Despite the problems just mentioned pectination interpreted

carefully has its value and it is not an arbitrary process As one reads the terminals of a

pectinate tree from left to right adjacent terminals will be separated by apomorphies

that will cumulate in an appropriate fashion When thinking of a book or a herbarium

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

23

sequence (see Haston et al 2007) this is of some value Nymphaeales and

Austrobaileyales are here adjacent on the tree but they could be separated by

hundreds of families in the sequence merely by rotating only the node from which

Nymphaeales and their sister group (all other angiosperms minus about 75 species)

arise If adjacent in a book or herbarium or as in these summary pages then it is

relatively easy to relate their characters both apomorphic and plesiomorphic but if

separated by hundreds of pages or two floors in a large building then it is less easy to

get anything from the sequence (Since all orders in the pages of the main part of this

site are preceded by the apomorphies of all nodes immediately below them in the seed

plant phylogeny and because of the linkages that have been built in to the site this

problem is much less serious there there is no sequence) So the particular sequence of

all seed plants that is based on a phylogenetic classification but is to be used in herbaria

or any other place where taxa have to be arranged linearly as in this part of APweb

will be one that maximizes the number of taxa that are both successive branches of the

tree and placed successively in the sequence Since specimens are generally filed under

families the outline of a new family sequence for arranging herbaria and books can now

be suggested (Haston et al 2007) and such a standardized sequence will further help

teaching and learning about plants

ON THINKING ABOUT APOMORPHIES

Identifying apomorphies is important because understanding the evolution of

morphology in the broad sense - ie including anatomy chemistry etc - in the context

of diversification is one of our major goals For identifying apomorphies several

preconditions must be met One needs to have an accurate well supported phylogeny

one has to have examined the right taxa both from the point of view of morphology and

molecules one has to have coded the characters right (ie delimited states

appropriately) and one has to use an appropriate model of evolution when fitting the

variation to the tree (see eg Omland 1999 Stevens 2006b) If the distributions of

apomorphies below are compared with those in other studies (eg Turgeon et al 2001

Bremer et al 2001 Endress 2001 Albach et al 2001a Judd amp Olmstead 2004 D Soltis

et al 2005b Zhang et al 2006 Judd et al 2007) the reader may find a number of

differences Although I have integrated such studies as far as is possible there are five

reasons that link to the preconditions just mentioned why there may be differences

where features are placed on trees

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

24

1 I may not have found all the information about a particular character there may be

disagreement over its interpretation or I have added information

2 The sampling of nearly all molecular studies is very incomplete (see Salisbury amp Kim

2001 for problems caused by sampling) indeed it is commonplace to decry the

incompleteness of molecular sampling But not only is the sampling in molecular studies

often less than we might wish that of the morphological and chemical characters whose

evolution we are interested in understanding is also often very poor So for many

anatomical chemical and embryological characters that are confidently said to

characterise families and other groups we all too often have no idea if those characters

are applicable to the whole clade or just to a subgroup within that clade However as

our sampling improves we can locate changes on the tree more precisely if in

unexpected positions Thus is is sometimes suggested that Ericaceae have ellagic acid

(Soltis et al 2005b) in fact the little evidence we have indicates that only a small clade

of some 80 species has ellagic acid and the rest of the family - itself alone about one

third of all Ericales - does not Similarly Ericaceae are noted for their

ectendomycorrhizal associations but Enkianthus sister to all other Ericaceae

apparently lacks these (Abe 2005) Thus these mycorrhizae and their associated

distinctive hair roots are unlikely to be apomorphies of Ericaceae - however it appears

that other families near Ericaceae may also have ectendomycorrhizae (see Asai 1934)

3 I am fitting characters to a very conservative tree with many polytomies although the

nodes that are utilised are for the most part strongly supported polytomies make the

optimisation of characters that is the assigment of character state change to a

particular node on the tree notably difficult (eg Madison amp Madison 2002) In nearly

all studies of the evolution of characters distributions of characters are optimised on a

more or less fully resolved tree and the construction of supertrees may yield yet more

detailed hypotheses of relationships (for literature on supertrees see Cotton amp

Wilkinson 2007 2008) Of course some nodes on such fully resolved trees andor

supertrees may have little support and optimisations of characters on such trees may

carry correspondingly little conviction Even parts of some of the trees used here have

poor support eg relationships within aquatic Alismatales etc although I indicate

when this is the case

4 Exactly how one goes about optimising a character on a tree is critically important

Even using simple parsimony optimisations (ACCTRAN or DELTRAN ACCelerated

TRANsitions or DELayed TRANsitions) the position of synapomorphies on trees - and

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

25

hence our ideas of evolution - may differ and this is still more true if one uses maximum

likelihood or Bayesian methods and makes apparently reasonable suggestions about

weighting gains over losses (eg Donoghue amp Ackerley 1996 Cunningham et al 1998

Omland 1997 1999 Ree amp Donoghue 1999 Polly 2001 Webster amp Purvis 2001

Ronquist 2004 Crisp amp Cook 2005) Sannier et al (2007) show how in in Arecaceae that

where on a phylogenetic tree one might peg changes in microsporogenesis will depend

on the methods one uses to do this and Pedersen et al (2007) discuss the sometimes

very substantial effect of node support on the posterior probabilities of ancestral

character states Here I use parsimony optimization not always as explicit as it might be

but I have often indicated where there are particularly important uncertainties as to the

positions of particular character changes on the tree

5 Finally although I have paid quite a lot of attention to the delimitation of the

character states that make up all the characterizations I have not spent enough time on

this critical operation If we are interested in understanding evolution then fitting the

basic variation - not character states - to a tree in principle allows greater flexibility in

understanding morphology in the context of local phylogenies (see also Stevens 2000

Endress 2005c) However many character states used here are delimited globally that

is they are circumscribed in the context of the variation shown by individual characters

across all angiosperms andor in the context of classic ideas of character evolution

Character states often have arbitrary limits and serve best to communicate

information whether they are in fact suitable for either phylogenetic analysis or

understanding evolution are separate issues Studies have rather unsurprisingly

perhaps but importantly shown that dividing the one character into different sets of

states may yield differing ideas of evolution of that character (eg Lamb Frye amp Kron

2003 Hibbett 2004) When looking at trees on which character states are optimised

one should bear in mind the problems surrounding the delimitation of states (eg

Stevens 2000 2006b) and the danger of using pollen or other types - constructs

based on many characters that vary independently but which effectively get lost in

these types thus Blackmore et al (2009) decomposed the pollen types in Asteraceae

into 52 characters

I use Remanes three main criteria of homology or better similarity when

determining the basic similarity of structures on different organisms (see Remane 1952)

These criteria are special properties position and intermediates Special properties

include anatomical or chemical characters gene expression data etc Position refers

to the position of an organ with respect to landmarks on the plant Although plants are

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

26

plants and landmarks are not as fixed as one might like trying to work out the relative

positions of parts is a good way of understanding morphology so long as one does not

try to out-do Procrustes Intermediates include intermediates found on the same

plant or on different plants Intermediates may observed during development or be

adult structures Thus one may find changes during development which suggest what

the nature of a particular structure is and very different stuctures on different plants

may turn out to be similar early in development Or one can simply compare different

taxa and see that structures that appear to be very different when comparing two

different taxa can be linked morphologically by looking at other taxa

Nevertheless the use of these criteria may not yield an unambiguous answer as to

what a structure is even given a solid phylogeny and an improved understanding of

development (see Jaramillo amp Kramer 2007 for a useful discussion) As Endress (2005c)

observed a number of features - position function development shape anatomy

histology gene activity and relationships to other taxa that clearly have petals - can be

used to distinguish a petal (for example) from other floral structures if a petal does not

have one of these features is it thereby not a petal Thus Maturen et al (2005) recently

found that floral organ diversity genes (B and C) were expressed in the large white

inflorescence bracts of Cornus (see also Costa et al 2005) Peney et al (2005) noted that

not all monosulcate pollen grains in monocots have the same developmental pathway

and that as a result such pollen might not have the same ancestral state Reeves and

Olmstead (2003) suggested that the genetic mechanisms causing monosymmetry in

Lamiales and Solanales were different and Serna and Martin (2006) described similar

problems with the development of hairs in Arabidopsis when compared with that of

hairs in Antirrhinum and Solanaceae Indeed as one perhaps might expect delimitation

of states and characters does not necessarily become easier with increasing knowledge

of development etc Thus Buzgo et al (2004) Matthews and Endress (2005) and others

have shown how hard it can be to distinguish between eg prophylls and other floral

structures as their behaviour is studied during the course of the development of the

flower Similarly at what concentration is a particular secondary metabolite deemed to

be present (Waterman (2007)

The validity of the approach used here that of fitting morphological variation to a

largely molecular-based tree may be questioned However I think it rather unlikely that

well-supported molecular branches will be overturned by morphological data Indeed

analyses of morphological data alone do provide support for many of the clades evident

in molecule-only analyses and analyses with morphological and molecular data

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

27

together may lead to increased support for clades (eg Hufford 1992 Nandi et al 1998

[but adding morphological data reduces support for a number of critical clades too]

Doyle amp Endress 2000 however in none of these papers is the use of morphology

without ambiguity) It is unfortunately clear that the use of morphology alone may not

suggest problems in the phylogenetic placement of taxa that later turn out to have been

wrongly included (eg Zhang et al 1992) Thus I have been wary of putting much weight

on clades that have only morphological support but note that I have treated molecule-

based clades with low bootstrap or jacknife support values (esp below 70) or low

posterior probabilies (below 095) likewise Although I may have been mistaken in

placing so much emphasis on molecular data in terms of providing the basic

phylogenetic framework for angiosperms morphological and molecular data are only

very rarely in irreconcilably strong conflict There are indeed a few places where the

conflict seems extreme These include the relative positions of the Monimiaceae and

Hernandiaceae (Laurales) the position of Hanguanaceae (Commelinales [as here] or

Zingiberales) and of Triplostegia (is it in Dipsacaceae or Valerianaceae - see

Dipsacales) Fossils are unlikely to affect the topologies of the trees presented here but

see below for their importance in understanding morphological evolution in general and

the evolution of angiosperms in particular Nevertheless some largely reject the idea

that trees based on molecular data alone can recover phylogenetic relationships

especially when branching points are old and prefer to used trees based on analysis of

morphological data including those taken from fossils (Hilton amp Bateman 2006 Farjon

2007)

All in all however the extent of the congruence between morphological and

molecular data is impressive and heartening and many clades can be characterised

morphologically It seemed in 1998 that there were no unambiguous morphological

synapomorphies for angiosperm orders (K Bremer 2000) and this is still true if by

unambiguous is meant non-homoplasious However many orders and other clades

have synapomorphies even if these may be indistinguishable at least at the current

level of morphological and developmental knowledge from parallel occurences

elsewhere As our morphological knowledge increases so too does the number of

apomorphies

Endress and Matthews (2006a) emphasize the importance of tendencies and

developmental constraints when thinking about characters of clades within the rosids -

many characters or character combinations occur in a rather sporadic fashion within

the clade and apparently notably less frequently outside it There are a number of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

28

examples of tendencies here and in the individual order pages such as the features

enclosed by parentheses in groups above the level of family eg see (cuticular waxes

as aggregated rodlets) for the commelinids and the discussion of the distribution of

polyandry that is flowers with numerous stamens in the asterid I + II groups etc

Tendencies may involve more than single characters When thinking of the

associations of features like integument thickness and vascularization of the integument

(Endress amp Matthews 2006a) one can imagine fairly simple developmental

preconditions being operative As Endress and Matthews (2006a) note it is difficult to

imagine vascular tissue developing in an integument only two cells thick Changes here

would be loosely correlated if morphologically linked However the strongly correlated

changes noted by Givnish et al (2005) are ecologically linked but are presumably

morphologicallydevelopmentally independent When there seem to be characters

evolving more or less together tests can be carried out to see if the changes are

concentrated on certain branches of the tree (eg Maddison 1990 Sanderson 1991

Maddison amp Maddison 2000) Such tests have rarely been carried out at higher levels in

angiosperms

One of the most striking examples of a tendency is the distribution of N-fixation

restricted as it is to a monophyletic group of four clades although it has arisen seven (or

perhaps several more) times independently within the N-fixing clade and several

members of two quite different kinds of bacteria are involved (eg Clawson et al 2004

Elliott et al 2007 Sprent amp James 2007) Other examples of work that bears on the issue

of tendencies include the findings that flowers of polysymmetrical Arabidopsis have

genes like TCP1 that are expressed asymmetrically during early development and TCP1

is a probable orthologue of the well-known CYC gene of Antirrhinum that is involved in

the development of monosymmetric slowers there (Cubas et al 2001 Costa et al 2005

etc) Parallelism might build on this underlying morphologically cryptic monosymmetry

even if details of the genetic mechanisms causing the monosymmetry evident in

particular groups may be different (Reeves amp Olmstead 2003 Cubas 2004 see above)

Similarly there have been several recent suggestions that the capability to synthesise a

particular metabolite may be switched off but not lost and so can sometimes be

reacquired (eg Wink amp Witte 1983 Wink 2003 Liscombe et al 2005 Larsson 2007

Waterman 2007) Hence perhaps the rather spotty distribution of many secondary

metabolites like ellagic acid the indole alkaloid camptothecin iridoids etc when

considered in the context of phylogenies As a non-botanical example - but a rather nice

one - Salwini-Plawen and Mayr (1961) suggested some time ago that there has been

considerable parallelism (40-65 or more independent origins) in the evolution of eyes in

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

29

metazoans However Pax 6 seems to be a master control gene that is involved in eye

formation perhaps throughout all bilateralians (eg Gehring amp Ikeo 1999 Erwin amp

Davidson 2002) which changes the nature of the problem Note however that in

general our invocation of evolutionary tendencies to explain the patterns of

distributions of characters is really an admission that we do not understand those

patterns

As we find out more about variation we will probably find fewer and fewer features

constant throughout a group Most unqualified statements of presence and absence

should properly be qualified as usually present or usually absent if one is thinking of

the characterisations on the order pages as encompassing the total variation within a

clade Thus Pistia alone among monocots as so far known has sieve tube plastids with

starch grains not protein crystals However this does not affect the fact that sieve tube

plastids with cuneate protein crystals and lacking starch grains are an apomorphy for

monocots Furthermore fossils in a number of cases suggest character combinations

unknown in extant taxa as may be seen in the discussions of Fagaceae Platanaceae

Iteaceae Calycanthaceae etc Confusing the issue there may be questions as to where

exactly on the tree a particular fossil is to be placed (eg see Nymphaeaceae

Calycanthaceae Archaefructus etc)

To summarize given our current understandings of both phylogenies and

characters evolution of some characters in which we are interested seems very labile

(see eg D Soltis et al 2005b Endress and Matthews 2006a Stevens 2006b Ekman et

al 2008 for a good recent study) and I have been cautious when talking about character

evolution Much effort must continue to be spent in summarizing characters of well-

established clades at all levels providing features by which they may be recognized and

signaling synapomorphies Remember that (1) the basic morphological anatomical and

chemical knowledge of many critical taxa is woefully incomplete (2) different

assumptions about character evolution may greatly affect the position of

synapomorphies on trees (3) in many cases relationships within and between many

groups are too uncertain at present to worry very much about synapomorphies and

(4) we must be clear about what we do and do not not know As mentioned above I

have indicated in a number of places where there are particularly important

uncertainties as to where characters should be placed on the tree Nevertheless it is a

relatively easy matter to update notes such as these and it can be a simple matter to

incorporate new data on characters that have never before been considered in the

context of a tree Much basic - and unfortunately perhaps unfashionable - work must be

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

30

carried to clarify the distribution of morphological anatomical and chemical characters

for examples of what can to be done see the work of P K Endress and collaborators

(floral morphology and development) and S R Jensen and collaborators (iridoids) But

acquiring information about nodal anatomy cell and tissue distribution nodal anatomy

and the like is not difficult at all and should be generally encouraged

But all this is merely a necesary prelude to the understanding of evolution For this

one needs to know a lot more including dating the phylogeny understanding the

function(s) of characters factoring in relevant aspects of the palaeoenvironment in

which an apomorphy first appeared etc

SUMMARY OF THE SYSTEM

Below is a formalised summary of the relationships within orders of the families of seed

plants There are a few families that are not recognised even as options in APG II and

vice versa as well as a few extra orders The families to be recognised in the most

recent edition of Mabberleys The Plant Book (Mabberley 2008) are also largely

consistent with those below But all differences are trivial and will - I hope - eventually

disappear

Square brackets - [] - enclose clades the plus sign - + - designates sister taxa a

comma - - denotes part of a polytomy and quotation marks - - denotes a

paraphyletic group

SEED PLANTS

GYMNOSPERMS

Cycadales

Cycadaceae + Zamiaceae

Ginkgoales

Ginkgoaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

31

Pinales

Pinaceae [[Araucariaceae [Podocarpaceae + Phyllocladaceae]] [Sciadopityaceae

[Taxaceae + Cupressaceae]]]

Gnetales

Ephedraceae [Gnetaceae + Welwitschiaceae]

ANGIOSPERMSFLOWERING PLANTS

Amborellales

Amborellaceae

Nymphaeales

Hydatellaceae [Cabombaceae + Nymphaeaceae]

Austrobaileyales

Austrobaileyaceae [Schisandraceae + Trimeniaceae]

Chloranthales

Chloranthaceae

MAGNOLIIDS

Magnoliales

Myristicaceae [Magnoliaceae [[Himantandraceae + Degeneriaceae] [Eupomatiaceae +

Annonaceae]]]

Laurales

Calycanthaceae [[Siparunaceae [Gomortegaceae + Atherospermataceae]] [Monimiaceae

[Hernandiaceae + Lauraceae]]]

Canellales

Canellaceae + Winteraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

32

Piperales

[Hydnoraceae + Aristolochiaceae] [Piperaceae + Saururaceae]

MONOCOTS

Acorales

Acoraceae

Alismatales

Araceae [Tofieldiaceae [[Alismataceae [Hydrocharitaceae + Butomaceae]]

[Scheuchzeriaceae [Aponogetonaceae [Juncaginaceae [Maundiaceae [[Posidoniaceae

[Ruppiaceae + Cymodoceaceae]] [Zosteraceae + Potamogetonaceae]]]]]]]

Petrosaviales

Petrosaviaceae

Dioscoreales

Nartheciaceae [[Taccaceae + Thismiaceae] [Burmanniaceae + Dioscoreaceae]]

Pandanales

Velloziaceae Triuridaceae Stemonaceae [Pandanaceae + Cyclanthaceae]

Liliales

Corsiaceae [Campynemataceae [Petermanniaceae [Colchicaceae + Alstroemeriaceae]]

Melanthiaceae [[Philesiaceae + Rhipogonaceae] [Smilacaceae + Liliaceae]]]

Asparagales

Orchidaceae [[Boryaceae [Blandfordiaceae [Lanariaceae [Asteliaceae + Hypoxidaceae]]]]

[[Ixioliriaceae + Tecophilaeaceae] [Doryanthaceae [Iridaceae [Xeronemataceae

[Xanthorrhoeaceae [Alliaceae + Asparagaceae]]]]]]]

COMMELINIDS

Unplaced

Dasypogonaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

33

Arecales

Arecaceae

Poales

[Typhaceae + Bromeliaceae] [Rapateaceae [[Thurniaceae [Juncaceae + Cyperaceae]]

[[Mayacaceae [Eriocaulaceae + Xyridaceae]] [Flagellariaceae [Anarthriaceae

[Centrolepidaceae + Restionaceae]]] [Joinvilleaceae [Ecdeiocoleaceae + Poaceae]]]]]]]

Commelinales

[Commelinaceae + Hanguanaceae] [Philydraceae [Haemodoraceae + Pontederiaceae]]

Zingiberales

Musaceae [Strelitziaceae + Lowiaceae] Heliconiaceae [[Cannaceae + Marantaceae]

[Costaceae + Zingiberaceae]]

Ceratophyllales

Ceratophyllaceae

EUDICOTS

Ranunculales

Eupteleaceae [Papaveraceae [[[Lardizabalaceae + Circaeasteraceae] [Menispermaceae

[Berberidaceae + Ranunculaceae]]]]

Sabiales

Sabiaceae

Proteales

Nelumbonaceae [Platanaceae + Proteaceae]

Trochodendrales

Trochodendraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

34

Buxales

Haptanthaceae [Buxaceae + Didymelaceae]

CORE EUDICOTS

Gunnerales

Gunneraceae + Myrothamnaceae

Dilleniales

Dilleniaceae

Saxifragales

Peridiscaceae [[Paeoniaceae [Altingiaceae [Hamamelidaceae [Cercidiphyllaceae +

Daphniphyllaceae]]]] [[Crassulaceae [Aphanopetalaceae [Tetracarpaeaceae

[Penthoraceae + Haloragaceae]]]] [Iteaceae [Grossulariaceae + Saxifragaceae]]]]

Cynomoriaceae unplaced

Vitales

Vitaceae

ROSIDS

FABIDROSID I

Zygophyllales

Krameriaceae + Zygophyllaceae

Celastrales

Lepidobotryaceae + Celastraceae

Oxalidales

Huaceae [[Connaraceae + Oxalidaceae] [Cunoniaceae [Elaeocarpaceae [Brunelliaceae +

Cephalotaceae]]]]

Malpighiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

35

[Achariaceae [Goupiaceae [Violaceae + Passifloraceae] [Lacistemataceae Salicaceae]]

[Lophopyxidaceae Putranjivaceae] [Ctenolophonaceae [Erythroxylaceae +

Rhizophoraceae]] Linaceae Ixonanthaceae Humiriaceae Irvingiaceae

Centroplacaceae Pandaceae [[Bonnetiaceae + Clusiaceae] [Calophyllaceae

[Hypericaceae + Podostemaceae]]] [Malpighiaceae + Elatinaceae] Ochnaceae

[Peraceae [Rafflesiaceae + Euphorbiaceae]] [Phyllanthaceae + Picrodendraceae]

[Balanopaceae [[Trigoniaceae + Dichapetalaceae] [Chrysobalanaceae +

Euphroniaceae]]] Caryocaraceae

N-FIXING CLADE

Fabales

Quillajaceae [Fabaceae [Polygalaceae + Surianaceae]]

Rosales

Rosaceae [[Barbeyaceae [Elaeagnaceae Dirachmaceae Rhamnaceae]] [Ulmaceae

[Cannabaceae [Moraceae + Urticaceae]]]]

Cucurbitales

Anisophylleaceae [[Corynocarpaceae + Coriariaceae] [Cucurbitaceae [Tetramelaceae

[Datiscaceae + Begoniaceae]]]] Apodanthaceae

Fagales

Nothofagaceae [Fagaceae [[Myricaceae + Juglandaceae] [Casuarinaceae

[Ticodendraceae + Betulaceae]]]]

MALVIDROSID II

pgtGeraniales

Geraniaceae [[Melianthaceae + Francoaceae] [Vivianaceae + Ledocarpaceae]]

Myrtales

Combretaceae [[Onagraceae + Lythraceae] [[Vochysiaceae + Myrtaceae]

[Melastomataceae [Crypteroniaceae [Alzateaceae + Penaeaceae]]]]]

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

36

Crossosomatales

[Staphyleaceae [Guamatelaceae [Crossosomataceae + Stachyuraceae]]] [Aphloiaceae

[Geissolomataceae + Strasburgeriaceae]]

Picramniales

Picramniaceae

Sapindales

Biebersteiniaceae Nitrariaceae [Kirkiaceae [Anacardiaceae + Burseraceae]]

[Sapindaceae [Simaroubaceae Rutaceae Meliaceae]]

Huerteales

Gerradinaceae [Dipentodontaceae + Tapisciaceae]

Malvales

Neuradaceae [Thymelaeaceae [Sphaerosepalaceae Bixaceae [Cistaceae

[Sarcolaenaceae + Dipterocarpaceae]] [Cytinaceae + Muntingiaceae] Malvaceae]]

Brassicales

[Akaniaceae + Tropaeolaceae] [[Moringaceae + Caricaceae] [Setchellanthaceae

[Limnanthaceae [[Koeberliniaceae [Bataceae + Salvadoraceae]] [Emblingiaceae

[Pentadiplandraceae [Gyrostemonaceae Resedaceae Stixis and relatives] Tovariaceae

[Capparaceae [Cleomaceae + Brassicaceae]]]]]]]]

Berberidopsidales

Aextoxicaceae + Berberidopsidaceae

Santalales

Erythropalaceae Olacaceae [[Loranthaceae [Misodendraceae + Schoepfiaceae]]

[Opiliaceae + Santalaceae]] Balanophoraceae unplaced

Caryophyllales

[[Droseraceae [Nepenthaceae [Drosophyllaceae [Ancistrocladaceae +

Dioncophylleaceae]]]]] [[Frankeniaceae + Tamaricaceae] [Polygonaceae +

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

37

Plumbaginaceae]]] [Rhabdodendraceae [Simmondsiaceae [[Asteropeiaceae +

Physenaceae] [[Caryophyllaceae [Achatocarpaceae + Amaranthaceae]]

[Stegnospermataceae [Limeaceae [[Lophocarpaceae [Barbeuiaceae [Aizoaceae

[Sarcobataceae Phytolaccaceae Nyctaginaceae]]]] [Molluginaceae [Halophytaceae

Montiaceae Basellaceae Didiereaceae [Talinaceae [Portulacaceae [Anacampseros etc +

Cactaceae]]]]]]]]]]]]]

ASTERIDS

Cornales

[Cornaceae + Nyssaceae] [Hydrangeaceae + Loasaceae] Hydrostachyaceae [Curtisiaceae

+ Grubbiaceae]

Ericales

[Balsaminaceae [Marcgraviaceae + Tetrameristaceae]] [[Polemoniaceae +

Fouquieraceae] Lecythidaceae [[Sladeniaceae + Pentaphylacaceae] [Sapotaceae

[Ebenaceae [Maesaceae [Theophrastaceae [Myrsinaceae + Primulaceae]]]]]

[Mitrastemonaceae Theaceae [Symplocaceae [Styracaceae + Diapensiaceae]]

[[Sarraceniaceae [Roridulaceae + Actinidiaceae] [Clethraceae [Cyrillaceae +

Ericaceae]]]]]]

LAMIIDASTERID I

Unplaced

Metteniusaceae Oncothecaceae Icacinaceae and some unplaced ex-Icacinaceae

Garryales

Garryaceae + Eucommiaceae

Unplaced

Boraginaceae Vahliaceae

Gentianales

Rubiaceae [Gentianaceae [Loganiaceae [Gelsemiaceae + Apocynaceae]]]

Lamiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

38

Plocospermataceae [Oleaceae [Tetrachondraceae [Calceolariaceae

[Gesneriaceae Peltanthera [Pedaliaceae Carlemanniaceae Martyniaceae Stilbaceae

Plantaginaceae Linderniaceae Bignoniaceae Verbenaceae Lamiaceae Paulowniaceae

Schlegeliaceae Thomandersiaceae Phrymaceae Rehmannia Acanthaceae

Scrophulariaceae Orobanchaceae Byblidaceae Lentibulariaceae]]]]]

Solanales

[Montiniaceae [Sphenocleaceae + Hydroleaceae]] [Convolvulaceae + Solanaceae]

CAMPANULIDASTERID II

Aquifoliales

[[Cardiopteridaceae + Stemonuraceae] [Aquifoliaceae [Helwingiaceae +

Phyllonomaceae]]

Asterales

[Rousseaceae + Campanulaceae] Pentaphragmataceae [[Alseuosmiaceae [Phellinaceae

+ Argophyllaceae]] Stylidiaceae [Menyanthaceae [Goodeniaceae [Calyceraceae +

Asteraceae]]]]

Escalloniales

Escalloniaceae

Bruniales

[Bruniaceae + Columelliaceae]

Apiales

Pennantiaceae [Torricelliaceae [Griseliniaceae [Pittosporaceae [Araliaceae

[Myodocarpaceae + Apiaceae]]]]]

Paracryphiales

Paracryphiaceae

Dipsacales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

39

Adoxaceae [Diervillaceae [Caprifoliaceae [Linnaeaceae [Morinaceae [Dipsacaceae +

Valerianaceae]]]]]

LINKS TO ORDERS AND FAMILIES

[Back to Top]

MAIN GROUPINGS

Asterids commelinids core eudicots asterid 1 asterid

2 eudicots gymnosperms Magnoliophyta monocots N-fixing claderosids seed plants

ALPHABETICAL LISTING OF ALL ORDINAL NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | L | M | N | O | P | Q | R | S | T | U | V | W | X | Z

Abietales Acanthales Acerales Acorales Actinidiales Actinostrobales Adoxales Aescul

ales Agavales Aizoales AkanialesAlismatales Alliales Alseuosmiales Alstroemeriales

Altingiales Amaranthales Amaryllidales Amborellales AmbrosialesAmmiales Amomal

es Ancistrocladales Anisophylleales Annonales Anthobolales Apiales Apocynales Apo

nogetonalesAquifoliales Arales Araliales Aralidiales Araucariales Arecales Aristolochi

ales Athrotaxidales Asarales AsclepiadalesAsparagales Asphodelales Asparagales Ast

eliales Atriplicales Aucubales Austrobaileyales Avenales

Balanitales Balanopales Balanophorales Balsaminales Barbeyales Barclayales Batales

Begoniales BerberidalesBerberidopsidales Betulales Biebersteiniales Bignoniales Bix

ales Boraginales Brassicales Brexiales Bromeliales BrunialesBrunoniales Burmannial

es Burserales Butomales Buxales Byblidales

Cactales Callitrichales Calycanthales Calycerales Campanulales Campynematales Can

ellales Cannales CapparalesCaprifoliales Cardiopteridales Carduales Caricales Carle

manniales Caryophyllales Cassiales Casuarinales CelastralesCentrolepidales Cephalot

ales Cephalotaxales Ceratophyllales Cercidiphyllales Chenopodiales Chironiales Chlor

anthalesChrysobalanales Cinchonales Circaeasterales Cistales Citrales Cocosales Col

chicales Columelliales CombretalesCommelinales Connarales Convolvulales Coriarial

es Cornales Corylales Corynocarpales Crassulales CrossosomatalesCucurbitales Cunn

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

40

inghamiales Cunoniales Cupressales Cycadales Cyclanthales Cymodoceales Cynarales

CynomorialesCyperales Cyrillales Cytinales

Daphnales Daphniphyllales Dasypogonales Datiscales Degeneriales Desfontainiales D

iapensiales Didymelales DillenialesDioales Dioncophyllales Dioscoreales Diospyrales

Dipentodontales Dipsacales Droserales

Ebenales Echiales Elaeagnales Elaeocarpales Elatinales Elodeales Emmotales Empetr

ales Ephedrales Ericales EriocaulalesErythropalaless Escalloniales Eucommiales Euph

orbiales Eupomatiales Eupteleales Euryalaless

Fabales Fagales Falcatifoliales Ficales Flacourtiales Flagellariales Fouquieriales Franc

oales Frangulales

Galiales Garryales Geissolomatales Gentianales Geraniales Gesneriales Ginkgoales G

laucidiales Globulariales GnetalesGoodeniales Greyiales Griseliniales Grossulariales

Grubbiales Gunnerales Gyrocarpales Gyrostemonales

Haemodorales Haloragales Hamamelidales Hanguanales Heisteriales Helleborales He

lwingiales Himantandrales HippuridalesHomaliales Hortensiales Huales Huerteales

Hydatellales Hydnorales Hydrangeales Hydrastidales HydrocharitalesHydropeltidales

Hydrostachyales Hypericales Hypoxidales

Icacinales Illiciales Iridales Irvingales Iteales Ixiales

Jasminales Juglandales Julianiales Juncaginales Juncales

Lacistematales Lactoridales Lamiales Lardizabalales Laurales Lecythidales Ledocarpal

es Leitneriales LentibularialesLigustrales Liliales Limnanthales Linales Liriales Loasal

es Lobeliales Loganiales Lonicerales Loranthales LowialesLythrales

Magnoliales Malpighiales Malvales Marathrales Marcgraviales Mayacales Medusagy

nales Medusandrales MelanthialesMelastomatales Meliales Melianthales Meliosmal

es Menispermales Menyanthales Metteniusales MitrastemonalesMiyoshiales Moni

miales Moringales Musales Myricales Myristicales Myrothamnales Myrsinales Myrta

les

Najadales Nandinales Narcissales Nartheciales Nelumbonales Nepenthales Neuradal

es Nitrariales Nolanales NothofagalesNyctaginales Nymphaeales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

41

Ochnales Oenotherales Olacales Oleales Onagrales Oncothecales Opuntiales Orchid

ales Oxalidales

Saxifragales Pandales Pandanales Papaverales Paracryphiales Parasitaxales Paridales

Parnassiales PassifloralesPenaeales Pennantiales Pentaphragmatales Peridiscales P

etiveriales Petrosaviales Phellinales Philydrales PhyllanthalesPhysenales Phytolaccal

es Picramniales Pinales Pinguiculales Piperales Pittosporales Plantaginales Platanale

sPlumbaginales Poales Podocarpales Podophyllales Podostemales Polemoniales Pol

ygalales Polygonales PontederialesPortulacales Posidoniales Potamogetonales Primu

lales Proteales

Quercales Quillajales Quintiniales

Rafflesiales Ranunculales Rapateales Resedales Restionales Rhabdodendrales Rhamn

ales Rhinanthales RhizophoralesRhodorales Rhoipteleales Roridulales Rosales Rouss

eales Rubiales Ruppiales Rutales

Sabiales Salicales Salvadorales Samolales Samydales Sanguisorbales Santalales Sapin

dales Sapotales SarracenialesSaxegotheales Saxifragales Scheuchzeriales Sciadopitya

les Scleranthales Scrophulariales Scyphostegiales SedalesSilenales Simmondsiales S

milacales Solanales Sphenocleales Sphenostemonales Stangeriales Stellariales Stemo

nalesStilbales Stylidiales Styracales Surianales

Taccales Tamales Tamaricales Taxales Taxodiales Tecophilaeales Terebinthales Tern

stroemiales Theales TheligonalesThymelaeales Tiliales Tofieldiales Torricelliales Tov

ariales Tribelales Trilliales Trimeniales Triuridales TrochodendralesTropaeolales Turn

erales Typhales

Ulmales Urticales

Vacciniales Vahliales Vallisneriales Velloziales Veratrales Verbenales Viburnales Vinc

ales Violales Viscales VitalesVochysiales

Welwitschiales Winterales

Xanthorrhoeales Ximeniales Xyridales

Zamiales Zingiberales Zosterales Zygophyllales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

42

ALPHABETICAL LISTING OF ALL FAMILY NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | K | L | M | N | O | P | Q | R | S | T | U | V | W | X | Y

|Z

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In some cases clicking on a family synonym will send you directly to the subfamily in

which the genus of the basionym is to be found

Abietaceae Abolbodaceae Abrophyllaceae Acalyphaceae Acanthaceae [Acanthochla

mydaceae] Acarnaceaee AceraceaeAchariaceae Achatocarpaceae Achradaceae Achy

ranthaceae Acmopylaceae Acoraceae Acristaceae ActaeaceaeActinidiaceae Actinota

ceae Amentotaxaceae Adenogrammaceae Adoxaceae Aegialitidaceae Aegicerataceae

Aegilopaceae Aeginetiaceaee Aegiphilaceae Aesculaceae Aextoxicaceae Agapanthac

eae Agavaceae AgdestidaceaeAgialidaceae Agrimoniaceae Agrostidaceae Ailanthace

ae Aitoniaceae Aizoaceae Akaniaceae Alangiaceae AlchemillaceaeAldrovandaceae A

lismataceae Alliaceae Allioniaceae Allophylaceae Aloaceae Alopecuraceae Alpiniacea

e AlseuosmiaceaeAlsinaceae Alsodeiaceae Alstroemeriaceae Altingiaceae Alzateacea

e Amaranthaceae Amaryllidaceae Amborellaceae Ambrosiaceae Amentotaxaceae A

mmanniaceae Ammiaceae Amomaceae Ampelidopsaceae Amygdalaceae Amyridacea

eAnacardiaceae Anagallidaceae Anarthriaceae Anchusaceae Ancistrocladaceae Andr

omedaceae AndropogonaceaeAndrostachyaceae Androsynaceae Anemarrhenaceae

Anemonaceae Angelicaceae Anisophylleaceae AnnonaceaeAnomochloaceae Anopter

aceae Anrederaceae Anthemidaceae Anthericaceae Anthobolaceae Antidesmataceae

Antirrhinaceae Antoniaceae Aparinaceae Aphanopetalaceae Aphloiaceae Aphyllanth

aceae Apiaceae ApocynaceaeApodanthaceae Aponogetonaceae Aporusaceae Apose

ridaceae Apostasiaceae Aptandraceae AquifoliaceaeAquilariaceae Aquilegiaceae A

raceae Aragoaceae Araliaceae Aralidiaceaee Araucariaceae Arbutaceae Arceuthidace

aeArceuthobiaceae Arctostaphylaceae Arctotidaceae Ardisiaceae Arecaceae Argoph

yllaceae Arisaraceae AristolochiaceaeAristoteliaceae Arjonaceae Armeriaceae Artem

isiaceae Arthrotaxidaceae Artocarpaceae Arundinaceae ArundinellaceaeAsaraceae A

sclepiadaceae Ascyraceaee Asiraceae Asparagaceae Asperulaceae Asphodelaceae As

pidistraceaeeAsteliaceae Asteraceae Asteranthaceae Asteropeiaceae Astragalaceae

Astrocarpaceae AthanasiaceaeAtherospermataceae Atriplicaceae Atropaceae Aucuba

ceae Aurantiaceae Austrobaileyaceae Austrotaxaceae AvenaceaeAverrhoaceae Avet

raceae Avicenniaceae Azaleaceae Azimaceae

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Balanitaceae Balanopaceae Balanophoraceae Balsameaceae Balsaminaceae Bambusa

ceae Barbaceniaceae BarbeuiaceaeBarbeyaceae Barclayaceae Barringtoniaceae Base

llaceae Bataceae Baueraceae Bauhiniaceae Baxteriaceae BegoniaceaeBehniaceae B

elangeraceae Belloniaceae Bembiciaceae Berberidaceae Berberidopsidaceae Berryac

eaee BersamaceaeBertyaceae Berzeliaceae Besleriaceae Betaceae Betulaceae Biebe

rsteiniaceae Bifariaceae Bignoniaceae BischofiaceaeBixaceae Blakeaceae Blandfordia

ceae Blattiaceae Blepharocaryaceae Blitaceae Blyxaceae Boerlagellaceae Bolivaracea

eBombacaceae Bonnetiaceae Bontiaceae Boopidaceae Boraginaceae Borassaceae B

oroniaceae BoryaceaeBotryodendraceae Bougainvilleaceae Boweniaceae Brassicacea

e Brachycaulaceae Bretschneideraceae BrexiaceaeBromeliaceae Brownlowiaceae Br

unelliaceae Bruniaceae Brunoniaceae Brunsvigiaceae Bryoniaceae BuchneraceaeBuci

daceae Buddlejaceae Buglossaceae Bulbocodiaceae Bumeliaceae Bupleuraceae Burc

hardiaceae BurmanniaceaeBurseraceae Butneriaceae Butomaceae Buxaceae Byblida

ceae Byttneriaceae

Cabombaceae Cacaoaceae Cactaceae Caesalpiniaceae Caladiaceae Calamaceae Calce

olariaceae CalectasiaceaeCalendulaceae Callaceae Callicomaceae Calligonaceae Calli

traceae Callitrichaceae Calochortaceae CalophyllaceaeCalthaceae Calycanthaceae Ca

lyceraceae Cambogiaceae Camelliaceae Campanulaceae CampynemataceaeCanacom

yricaceae Canellaceae Cannabaceae Cannaceae Canopodaceae Canotiaceae Cansjera

ceae CapparaceaeCaprariaceae Caprifoliaceae Cardiopteridaceae Carduaceae Carica

ceae Carissaceae Carlemanniaceae CarpinaceaeCarpodetaceae Cartonemataceae Ca

ryocaraceae Caryophyllaceae Casearieaceae Cassiaceae CassipoureaceaeCassythacea

e Castaneaceae Castelaceae Casuarinaceae Catesbaeaceae Cathedraceae Cecropiace

ae CedraceaeCedrelaceae Celastraceae Celosiaceae Celtidaceae Centaureaceae Cen

trolepidaceae Centroplacaceae CepaceaeCephalanthaceae Cephalotaceae Cephalota

xaceae Cerastiaceae Ceratoniaceae Ceratophyllaceae CerberaceaeCercidiphyllaceae

Cercocarpaceae Cercodiaceae Cereaceae Cerinthaceae Ceroxylaceae Cestraceae Cev

alliaceaeChailletiaceae Chamaedoraceae Chamaemoraceae Chamelauciaceae Chaun

ochitonaceae Chelidoniaceae ChelonaceaeChenopodiaceae Chimonanthaceae Chingi

thamnaceae Chionographidaceae Chiranthodendraceae ChironiaceaeChlaenaceae Ch

loanthaceae Chloranthaceae Chloridaceae Chlorogalaceae Chrysobalanaceae Cicerac

eae CichoriaceaeCimicifugaceae Cinchoniaceae Circaeaceae Circaeasteraceae Cissac

eae Cistaceae Citraceae Clematidaceae CleomaceaeClethraceae Clusiaceae Cneorac

eae Cnestidaceae Cnicaceae Cobaeaceae Cochlospermaceae Cocosaceae Coffeaceae

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Colchicaceae Coleogynaceae Columelliaceae Combretaceae Commelinaceae Comocl

adiaceae Compositae CompsoaceaeConantheraceae Connaraceae Conostylidaceae C

onvallariaceae Convolvulaceae Cordiaceae CoreopsidaceaeCoriandraceae Coriariacea

e Coridaceae Corispermaceae Cornaceae Corokiaceae Coronillaceae Correaceae Cor

rigiolaceaeCorsiaceae Corydalaceae Corylaceae Corynocarpaceae Coryphaceae Cost

aceae Cotyledonaceae CoulaceaeCoutariaceae Coutoubiaceae Crassulaceae Crescen

tiaceae Cressaceae Crinaceae Crocaceae CroomiaceaeCrossosomataceae Crotonacea

e Cruciferae Cryptaceae Crypteroniaceae Cryptocorynaceae CryptomeriaceaeCtenol

ophonaceae Cucurbitaceae Cunninghamiaceae Cunoniaceae Cupressaceae Curcumac

eae Curtisiaceae CuscutaceaeCyananthaceae Cyanastraceae Cynanchaceae Cyanella

ceae Cycadaceae Cyclanthaceae CyclantheraceaeCyclocheilaceae Cydoniaceae Cymo

doceaceae Cynaraceae Cynocrambaceae Cynomoriaceae Cyperaceae CyphiaceaeCyp

hocarpaceae Cypripediaceae Cyrillaceae Cyrtandraceae Cyrtanthaceae Cytinaceae

Dacrycarpaceae Dactylanthaceae Damasoniaceae Daphnaceae Daphniphyllaceae Das

ypogonaceae DatiscaceaeDaturaceae Daucaceae Davidiaceae Davidsoniaceae Decais

neaceae Deeringiaceae Degeneriaceae DelphiniaceaeDendrophthoaceae Columelliac

eae Detariaceae Dialypetalanthaceae Dianellaceae Dianthaceae DiapensiaceaeDicha

petalaceae Dichondraceae Diclidantheraceae Dicrastylidiaceae Dictamnaceae Didiere

aceae DidymelaceaeDidymocarpaceae Diegodendraceae Diervillaceae Digitalidaceae

e Dilatridaceae Dilleniaceae Dioaceae DionaeaceaeDioncophyllaceae Dioscoreaceae

Diosmaceae Diospyraceae Dipentodontaceae Diphylleiaceae Diplolaenaceae Dipsaca

ceaeDipterocarpaceae Dirachmaceae Disanthaceae Diselmaceae Dodonaeaceae Do

mbeyaceae Donatiaceae DorsteniaceaeDortmannaceae Doryanthaceae Drabaceae D

racaenaceae Dracontiaceae Drimyidaceae Droseraceae DrosophyllaceaeDryadaceae

Duabangaceae Duckeodendraceae Dulongiaceae Durantaceae Durionaceae Dysphani

aceae

Ebenaceae Eccremidaceae Ecdeiocoleaceae Echiaceae Echinopaceae Ehretiaceae Ela

eagnaceae ElaeocarpaceaeElegiaceae Elatinaceae Ellisiaceae Ellisiophyllaceae Elodea

ceae Elytranthaceae Embeliaceae Emblingiaceae EmmotaceaeEmpetraceae Encepha

lartaceae Engelhardtiaceae Enhalaceae Epacridaceae Ephedraceae Ephemeranthacea

eEpilobiaceae Epimediaceae Eragrostidaceae Eremolepidaceae Eremosynaceae Erica

ceae Erinaceae EriocaulaceaeEriogonaceae Eriospermaceae Erodiaceae Erycibaceae

Eryngiaceae Erysimaceae Erythroniaceae ErythropalaceaeErythrospermaceae Erythro

xylaceae Escalloniaceae Eschscholziaceae Eucomidaceae Eucommiaceae Eucryphiace

aeEuonymaceae Eupatoriaceae Euphorbiaceae Euphrasiaceae Euphroniaceae Eupo

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matiaceae Eupteleaceae EuryalaceaeEustrephaceae Euthemidaceae Eutocaceae Exb

ucklandiaceae Exocarpaceae

Fabaceae Fagaceae Falcatifoliaceae Ferulaceae Festucaceae Ficaceae Ficoidaceae Fit

zroyaceae FlacourtiaceaeFlagellariaceae Flindersiaceae Foetidiaceae Forestieraceae

Fothergillaceae Fouquieriaceae Fragariaceae FrancoaceaeFrangulaceae Frankeniacea

e Fraxinaceae Fraxinellaceae Freycinetiaceae Fritillariaceae Fuchsiaceae Fumariacea

eFunkiaceae

Gaiadendraceae Galacaceae Galanthaceae Galaxiaceae Galedupaceae Galeniaceae G

aliaceae Garciniaceae GardeniaceaeGardneriaceae Garryaceae Geissolomataceae Ge

itonoplesiaceae Gelsemiaceae Geniostomaceae GentianaceaeGeonomataceae Geosir

idaceae Geraniaceae Gerrardinaceae Gesneriaceae Gethyllidaceae Gilliesiaceae Gina

lloaceaeGinkgoaceae Gisekiaceae Gladiolaceae Glaucidiaceae Glechomaceae Glinace

ae Globulariaceae Gnaphaliaceae GnetaceaeGoetzeaceae Gomortegaceae Gomphiac

eae Gomphrenaceae Gonystylaceae Goodeniaceae Gordoniaceae GouaniaceaeGoup

iaceae Gramineae Gratiolaceae Grewiaceae Greyiaceae Grielaceae Griseliniaceae Gr

onoviaceae GrossulariaceaeGrubbiaceae Guaiacanaceae Guamatelaceae Guettardac

eae Gunneraceae Gustaviaceae Guttiferae GyrocarpaceaeGyrostemonaceae

Hachetteaceae Haemanthaceae Haemodoraceae Halesiaceae Hallieraceae Halocarpa

ceae Halophilaceae HalophytaceaeHaloragaceae Hamamelidaceae Hanguanaceae H

aptanthaceae Harmandaceae Hebenstretiaceae HectorellaceaeHederaceae Hedyosm

aceae Hedyotidaceae Hedysaraceae Heisteriaceae Heleniaceae Heliamphoraceae H

elianthaceaeHelianthemaceae Helichrysaceae Heliconiaceae Helicteraceae Heliotropi

aceae Helleboraceae Heloniadaceae HelosidaceaeHelwingiaceae Hemerocallidaceae

Hemimeridaceae Hemiphylacaceae Henriqueziaceae Henslowiaceae Hermanniaceae

Hernandiaceae Herniariaceae Herreriaceae Hesperocallidaceae Heterantheraceae He

teropyxidaceae HeterostylaceaeHibbertiaceae Hibiscaceae Hilleriaceae Himantandra

ceae Hippocastanaceae Hippocrateaceae HippomanaceaeHippophaeaceae Hippurida

ceae Hirtellaceae Holacanthaceae Homaliaceae Hopkinsiaceae Hoplestigmataceae H

ordeaceaeHornschuchiaceae Hortensiaceae Hortoniaceae Hostaceae Houstoniaceae

Huaceae Huerteaceae HugoniaceaeHumbertiaceae Humiriaceae Hyacinthaceae Hyd

atellaceae Hydnoraceae Hydrangeaceae Hydrastidaceae HydrillaceaeHydrocharitacea

e Hydrocotylaceae Hydrogetonaceae Hydroleaceae Hydropeltidaceae Hydrophylacac

eae HydrophyllaceaeHydrostachyaceae Hymenocardiaceae Hypecoaceae Hyperanth

eraceae Hypericaceae Hypopityaceae HypoxidaceaeHypseocharitaceae

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Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta

ceae ImperatoriaceaeInocarpaceae Inulaceae Ionidiaceae Iriarteaceae Iridaceae Irvi

ngiaceae Isnardiaceae Isophysidaceae IteaceaeStrasburgeriaceae Ixiaceae Ixioliriace

ae Ixonanthaceae

Jalapaceae Jamboliferaceae Japonoliriaceae Jasionaceae Jasminiaceae Johnsoniaceae

Joinvilleaceae JuglandaceaeJulianiaceae Juncaceae Juncaginaceae Juniperaceae Jus

siaeaceae Justiciaceae

Kadsuraceae Kaliphoraceae Kalmiaceae Kaniaceae Kiggelariaceae Kingdoniaceae Kin

giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae

Krameriaceae

Labiatae Lacandoniaceae Lachenaliaceae Lacistemataceae Lactoridaceae Lactucaceae

Lagerstroemiaceaee LamiaceaeLampsanaceae Lanariaceae Langsdorffiaceae Lantan

aceae Lapageriaceae Lardizabalaceae Lasiopetalaceae LauraceaeLawsoniaceae Laxm

anniaceae Lecythidaceae Ledaceae Ledocarpaceae Leeaceae Leguminosae Leitneriac

eae LemnaceaeLennoaceae Lentibulariaceae Lentiscaceae Leoniaceae Leonticaceae

Lepidobotryaceae LepidocarpaceaeLepidocaryaceae Lepidocerataceae Leptaulaceae

Leptospermaceae Lepturaceae Lepuropetalaceae LeuctenbergiaceaeLibocedraceae

Licaniaceae Ligustraceae Lilaeaceae Liliaceae Limeaceae Limnanthaceae Limnocharit

aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace

ae Linderniaceae Lindleyaceae Linnaeaceae LippayaceaeLiriaceae Liriodendraceae Li

ssocarpaceae Littorellaceae Loasaceae Lobeliaceaee Loganiaceae LomandraceaeLoni

ceraceae Lopeziaceae Lophiocarpaceae Lophiolaceae Lophiraceae Lophophytaceae L

ophopyxidaceae LoranthaceaeLotaceae Lowiaceae Lupulaceaee Luxemburgiaceae Lu

zuriagaceae Lychnidaceae Lyciaceae LyginiaceaeLygodisodeaceae Lysimachiaceae Ly

thraceae

Macarisiaceae Mackinlayaceae Madiaceae Maesaceae Magnoliaceae Malaceae Male

sherbiaceae MalortieaceaeMalpighiaceae Malvaceae Mangiaceae Manicariaceae Ma

paniaceae Marantaceae Marathraceae MarcgraviaceaeMartyniaceae Mastixiaceae

Matricariaceae Maundiaceae Mayacaceae Medeolaceae Medusagynaceae Peridiscac

eaeMelaleucaceae Melampyraceae Melanophyllaceae Melanthiaceae Melastomatac

eae Meliaceae Melianthaceae MelicaceaeMeliosmaceae Melittidaceae Melochiacea

e Memecylaceae Mendonciaceae Menispermaceae Menthaceae MenyanthaceaeMe

nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

47

Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

Nypaceae Nyssaceae

Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

ceae Papilionaceae Pappophoraceae PapyraceaeParacryphiaceae Parasitaxaceae Par

ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

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hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

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cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

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ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 4: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

4

flagged as such and also monophyletic but a genus can never include families Generic

family etc names are simply words we use to denote appropriate parts of phylogenies

and minimal aspects of their relationships and the irreducible aspect of relationships

that is emphasized here is monophyly A monophyletic group is one containing only and

all descendents of a common ancestor and it is often characterizable by apomorphies

that is shared derived characters that appeared in the ancestral lineage or stem clade

of that monophyletic group

Thus I am using a flagged ie ranked hierarchy for naming taxa rather than the

unranked systems that have recently been proposed (see below) The rank terminations

used (-ales -aceae etc - the flags) merely suggest relative positions of groups in the

local hierarchy If Ericaceae and Vaccinioideae are part of the same monophyletic group

the latter must refer to a clade nested within the former even if neither can necessarily

be directly compared with Polemoniaceae and Cobaeoideae (other than all being

putatively monophyletic groups) Such a flagged hierarchy is useful as a mnemonic and

communication device (eg Stevens 2006a) It improves memorization and emphasis on

families and orders as here is a didactic device - families are monophyletic units useful

in communication major units learned by biologists and others world-wide

The distinction between grouping and ranking is extremely important as is how we

interpret the latter We can both agree that there is a genus Acer yet disagree as to

whether it should be in Aceraceae or submerged in Sapindaceae Although from one

point of view this disagreement is utterly trivial it can have profound consequences if

we misunderstand the nature of the classificatory hierarchy Taxa at the same rank are

equivalent only by designation and have nothing necessarily in common (unless they

are sister taxa) other than their monophyly Rank as used here has no meaning other

than signifying a monophyletic group that includes other monophyletic groups with

appropriately subordinate rank terminations Taxa at the same rank have often been

treated incorrectly as if they were equivalent by biologists attempting to understand

evolutionary or biogeographic problems (see Bertrand et al 2006 for detailed

discussion) even if those constructing or using classifications - including Darwin (1859) -

have been explicit about the non-equivalence of taxa at the one rank (Stevens 1997) In

fact rank terminations have relatively infrequently been used by taxonomists to reflect

absolute rank although Linnaeus (at least in theory) at the level of genus and species

may be such an example (Classifications where rank is absolute taxa at the same rank

somehow being comparable entities are class hierarchies in the strict sense - Stevens

2002 2006a) It has also been suggested that taxon rank be adjusted so that rank

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

5

somehow reflects the degree of morphological differences between taxa or that taxa at

the same rank be based on similar characters or show a similar amount of distinctness

This might be possible using phenetic methods of analysis but is very difficult if ones

classification is phylogeny-based as here it could promote instability if used in taxa

where such a criterion had not previously been used and it might also inadvertently

suggest that taxa might be equivalent (for an example see Fritsch et al 2008) There

have also been proposals that rank could reflect the age of the clade (eg Hennig 1966)

with clades that have diverged by a particular time all being given the same rank Apart

from the fact that aging times of divergence of clades is still a difficult enterprise huge

disruptions to our nomenclature would result Recent suggestions which invoke the use

of age in classifications focus on providing a standardized timeclip ie a set of letters

referring to a particular geological period that could simply be added to a conventional

taxon name (Avise amp Mitchell 2007) However even such timeclips are unlikely to come

into general use soon

It would be impossible even to think about a higher-level classification such as this

without the advances in our understanding of relationships made by the phylogenetic

analyses of molecular data carried out over the last twenty years One can then

integrate the data to be found in both classical and recent morphological studies with

these phylogenies For the dramatic changes in this area see for instance the

pessimistic attitude towards orders in Davis and Heywood (1963 107-108) The most

unsatisfactory taxon in Angiosperm classification they were indefinable their

circumscription was not fixed etc Families they thought were likely to be the largest

natural unit within the mono- or dicotyledons Along the same lines almost three

quarters of the orders (4459 monofamilial orders ignored) recognised by Cronquist

(1981) are not monophyletic ie they do not contain all and only the descendents of a

common ancestor Most of those orders that are monophyletic are very small

(Zingiberales with eight families are the largest) for families on the other hand about

two thirds (189273) are monophyletic

Turning now to phylogenetic classifications and the particular classification used

here Backlund and Bremer (1998) provide a useful discussion on the principles of

phylogenetic classification that is applicable at all levels apart from species (see also

Stevens 1998 also Albach et al 2004 Entwisle amp Weston 2005 Pfeil amp Crisp 2005 etc

for examples) Backlund and Bremers main principle is that taxa that are recognised

formally should be monophyletic However this does not indicate which particular

clades we might wish to name as families genera etc and talk about in general

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

6

conversation If a well-supported hypothesis of monophyly is a necessary prerequisite

for a group to be named it is not a sufficient prerequisite (but cf the PhyloCode -

Cantino amp de Queiroz 2006 Cantino et al 2007) Not all clades need be named indeed

it would barely be practicable (or practical) to do this To decide which clades should be

named additional criteria can be invoked Other things being equal it is helpful if 1 taxa

formally recognised are easily recognizable 2 groups that are well-established in the

literature are preserved 3 the size of groups is taken into account and 4

nomenclatural changes are minimized (Backlund amp Bremer 1998) Thus numerous small

groups have little to recommend them since individually they summarize little

information and tend to clog the memory while groups that are too big may be

amorphous Somewhat similarly Godfray and Knapp (2004 p 562) note that users

want stable informative and accessible classifications that enable easy identification

(see also Simon 2008) - although invoking users without specifying those who make up

this group is not very helpful This classification is for all interested in comparative

biology hence the emphasis on monophyly Although the clades named are sometimes

difficult to characterise there are as we shall see many ways of making such a

classification accessible to all

Problems with this emphasis on monophyly may be caused by reticulation events

such as hybridization endosymbiosis and lateral gene transfer but they are unlikely to

be common confusing factors here Genera can often be pegged to above the level at

which hybridization is at all common However in Poaceae-Pooideae-Triticeae there are

some intractable problems where extremely well established common usage and the

principle of monophyly are likely to remain at odds Many genera are certainly not

monophyletic here being allopolyploids and the genera are ultimately based on the

different genomes they contain (Dewey 1984 Loumlve 1984 Barkworth 2000 for a history

of Triticeae classification Petersen et al 2006) There is also extensive reticulation

reported within Danthonioideae (Pirie et al 2009) Evidence also increases of old

hybridization events elsewhere in flowering plants that at the very least cause

discordance between relationships suggested by different genomic compartments as in

Smedmark and Anderberg (2007 Sapotaceae) and Fehrer et al (2007) Morgan et al

(2009) and Pelser et al (2008 all Asteraceae - hybridization is likely to be quite a

problem at the generic level here) and genera like Medicagoare turning out to have

highly reticulating relationships at the species level (Maureira-Butler et al 2008) Of

course there are many problems trying to apply the concept of monophyly to species

and for many - but not all - biologists strict monophyly is a less important criterion at

this level (Funk amp Omland 2003 see much of the discussion in Houmlrandl 2006)

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

7

The major endosymbiotic events that characterize the clade of which flowering

plants are a part (and gave rise to chloroplasts and mitochondria) are very ancient and

cause no problems for the student of multicellular organisms However lateral gene

transfer has been detected in a number of situations between quite unrelated

organisms (eg Bergthorsson et al 2003 - Amborella and liverworts see also below but

cf Goremykin et al 2009) and it may be particularly common in mitochondria (Sanchez-

Puerta et al 2008) Here too there are no major problems providing one is careful

such transfers do however raise all sorts of interesting biological questions (see

Richardson amp Palmer 2007 for a summary) Although there is increasing evidence for the

importance of genome duplications - hybridization is one cause of this - at various times

during the evolution of seed plants and of palaeopolyploidy events within eg the

Lauraceae and Magnoliaceae clades (Soltis et al 2009 for a summary see eg

the Characters page for further discussion) these too do not currently seem to pose

problems for the adoption of monophyly as the sine qua non of groups to be recognised

formally in this phylogenetic classification but it can make detecting orthologous genes

difficult

The accessory principles of Backlund and Bremer (1998) should be used in

combination Thus keeping the monogenericPlatanaceae separate from its sister

taxon Proteaceae is justifiable Both are much-used names that signal well supported

well defined and easily recognisable groups that have long been recognised as distinct

have several synapomorphies and do indeed look very unlike each other Combining

the two would yield a clade with few obvious apomorphies not to mention the fact

that Nelumbonaceae should by the same logic (it is also monogeneric) also be included

in the expanded family On the other hand it is difficult to justify the continued

recognition of Callitrichaceae or Hippuridaceae monophyletic and distinctive although

they may be If they were recognised several poorly characterised clades would also

have to be carved out ofPlantaginaceae in any classification that aimed to convey a

comprehensive view of the worlds flora The continued recognition

of Valerianaceae and Dipsacaceae also tends to run into this problem (see also Pfeil amp

Crisp 2005 Orthia et al 2005 Albach 2008 etc for useful practical discussions of such

matters) But there are no absolute guidelines If Podostemaceae turn out to be sister

to Hypericaceae (for references here and elsewhere in the Introduction see the

individual families) the subsequent moderate dismemberment of Clusiaceae sl is not

be too high a price to pay for the continued recognition of Podostemaceae Hence the

somewhat provisional recognition of Hypericaceae and Calophyllaceae as well as

Clusiaceae below the families can all be recognized and the name Podostemaceae in

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

8

particular is very well established In a somewhat similar situation if Lemna and its

relatives are a clade sister to most other Araceae should they be recognised as a

separate family Gymnostachys a phenetically fairly distinctive taxon as well as the less

phenetically distinct Orontioideae would have to be recognised as a separate families

(or combined as a single family) too but Araceae in a somewhat restricted sense would

be somewhat more morphologically coherent although not greatly so and not notably

distinct However it is in the very nature of such decisions to be somewhat arbitrary

and unsatisfactory hence the emphasis on consensus classifications here and on

classifications as simply being a means to an end

A useful distinction can be drawn between crown groups and stem groups The

former are monophyletic and include the extant members of a clade and their

immediate common ancestor (see the figure below) The groups characterized in this

site are such groups Thus Proteaceae here are crown group Proteaceae apomorphies

like the single carpel four-merous perianth etc being found in this common ancestor

Stem groups on the other hand include all the members of a lineage immediately after

its split from its sister group and all branches of this lineage In the case of Proteaceae

the stem group would include everything after its split from its sister group

Platanaceae Obviously most of the organisms in the lineage that terminates in crown-

group Proteaceae are unknown only a few being known as fossils and it is not known

exactly where particular apomorphies of crown group Proteaceae evolved along this

lineage

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

9

Indeed there is no reason other than convention or convenience why any group

should not be segregated into several smaller monophyletic groups or merged to

produce a larger unit we can talk about one large thing or about several smaller things

Thinking about aspects of size findings in ethnobiology and cognitive psychology can be

used to suggest that a moderate number - probably fewer than 500 - of families is a

reasonable goal at which to aim and that groupings of taxa throughout any system

should be rather small in size (eg Berlin 1992 Stevens 1994 1997) Major systems such

as those of Linnaeus and Bentham and Hooker were constructed explicitly so as to ease

the burden on the memory (Stevens 1997 2002 see also Scharf 2007) the latter in

particular ensuring that all groups in their classification were relatively small often

containing three to eight immediately subordinate taxa - but by no means all their

groups were formally named Along the same lines Burtt (1977b) suggested that the

number of names at any rank should be at most one third those at the immediately

lower rank - and monotypic taxa might not need a formal name Consistent with such

ideas a fairly broad view of families and orders is taken here whenever the constraints

of monophyly and other criteria used when constructing classifications (see above)

permit The APG system is thus a convention consistent with ideas such as those

expressed by Backlund and Bremer (1998) for similar conventions see eg the Grass

Phylogeny Working Group (2001) and for fungi Hibbett et al (2007)

There are other views Thus Takhtajan (1997) has suggested that smaller families

are more natural This is incorrect Monophyletic groups that include fewer taxa -

Takhtajans smaller families - do not necessarily have more apomorphies than larger

groups even if members of smaller groups will always have more features in general in

common than the larger group that includes them That is they will have their

apomorphies their unique features as well as progressively more plesiomorphies

features found both in the small groups and in the larger clades of which they are a part

having more features in common is indeed one common meaning of more natural

(Note that the implication of the word natural has long been a group of the kind

[usually unspecified] that I think should be recognised and if other botanists had

different ideas too bad their groups were necessarily not natural [see Bather 1927] It

is thus rarely a helpful word and is not used here) Furthermore if this approach is

adopted we will find a slippery slope ahead By this kind of argument all families should

be very small since their members will have a great deal in common and so will be

most natural However as families (for example) are split the relationships that are

evident between the segregates and that were responsible for their being placed in a

single family in the first place will seem to necessitate the recognition of a new order

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

10

etc as is evident in Takhtajans own work - general taxonomic inflation is the result (see

also comparable suggestions in a cladistic context for Brassicales in particular - Ronse de

Craene amp Haston 2006) Such splitting is also questionable when teaching and learning

families since the student needs to understand the system as a whole However for

some genera removed from the families that until now have included them the

phenetic-classificatory-phylogenetic structure in their new home may mandate the

recognition of small families On the other hand Takhtajans suggestion that narrowly

defined families are more useful for phylogenetic studies may be true Indeed I have

more than once regretted prematurely combining groups whether species (in the

context of monographic work) or families (in the course of preparing these notes) but

this is largely a separate issue

Van Steenis (1978) Philipson (1987b) and others have rightly questioned the value

of splitting a group when ideas of the relationships of its constituent members have not

changed - that is very good reasons have to be provided for splitting a family if the

genera within it remain part of the same clade rather than belonging to another clade

Thus APG (2003) broadened the circumscription of Malvaceae because of the

parapolyphyly of some of the families that had historically been associated with it (Judd

amp Manchester 1997 Alverson et al 1999 Bayer et al 1999) These families particularly

Tiliaceae and Sterculiaceae were not at all easy to distinguish their close relationship

(see eg Brown 1814) and overall similarity had long been conceded and to some

workers at least their combination has come as something of a relief Although most of

the larger clades within Malvaceae sl remain difficult to distinguish even with flowers

Cheek (2007) opts for a wholesale and novel dismemberment into ten families

however the very good reasons for doing this are wanting

The same principles are of course applicable when it comes to dividing genera little

other than a headache is gained by splitting genera such

as Drosera and Gnetum (Doweld 2000) as has recently been proposed Thus if an

established genus divides into two (or more) clades this is not a signal for recognising

two groups at the same level - so here it could be argued that the dismemberment

of Pterostylis (Jones amp Clements 2002b) was somewhat unfortunate Along the same

lines if a newly-discovered taxon is sister to an existing named taxon - say a genus - this

does not necessitate the description of a separate genus for the newly described species

(cf Davis 2002) (Of course some [eg Thorne 1976] have suggested that the sizes of

gaps between groups at the same rank should be similar but any principle like this is

inherently flawed since morphological gaps are more unstable than phylogenetic

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

11

relationships and applying it across all flowering plants would both be difficult and cause

substantial changes - see also below) Humphreys and Linder (2009) provide a well-

documented survey of generic concepts in plants which the reader should consult they

note that generic limits (broad versus narrow) have oscillated historically and that

currently larger genera tend to be recognised because studies tend to be on a broader

scale than in the past

Note that invoking similarity or difference - whether qualified (considerable

similarities substantial differences) or not - in a cladistic context as justification for

combining or splitting taxa is not a particularly strong argument (see eg

CardiopteridaceaeStemonuraceae - Karingrehed 2002c) Similarity and difference can

neither be defined precisely since what may seem to be substantial similarities to me

may not to the next person nor are they likely to be stable in the face of our changing

knowledge of morphology and what might be synapomorphies

I might have prefered to merge some families recognised here or split others but

by and large I do not think my own preferences matter very much - and I take the same

position with regards to comparable preferences expressed by others Indeed the

bottom line is that in flagged hierarchies of the kind used here the limits of any

monophyletic unit generally taught and discussed particularly other than species can

be established only by convention and consensus (eg Stevens 2002 2006a Entwisle amp

Weston 2005) This is rather different from the reasonable observation that the

phylogeny itself cannot be achieved by consensus (Thorne 1976) for in most cases there

is a fact of the matter when it comes to relationships Given the increasing support for

the outlines of angiosperm phylogeny a stable consensus classification based on this

phylogeny seems attainable Indeed in addition to providing current ideas of

relationships of seed plants in a synthesised form this site is part of an attempt to build

such a consensus about the circumscription of taxa (see APG 1999 2003 2009 Grass

Phylogeny Working Group 2001 Mabberley 2008) Reaching such a consensus is vital

since what we know of angiosperm phylogeny allows a very large number of

classifications to be based on it and as we find out more the number of possible

classifications increases greatly Unfortunately however nature does not dictate what

the classification should be All classifications are constructed by humans to

communicate particular aspects of groups and relationships Our goals as systematists

are surely to produce robust hypotheses of relationships to understand the evolution of

morphology and the like - but not to argue ad nauseam whether something should be

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

12

a family or a subfamily That way surely lies madness and worse the discredit of our

discipline

There are similar issues whatever naming system is used Thus in phylogenetic

naming (Baum et al 1998 for an example but cf Baum et al 2004 for the PhyloCode

see Cantino amp De Queiroz 2006) an unflagged hierarchy is used in which any

terminations of names used are uninformative about the relative position of taxa If one

adopts the principle of phylogenetic naming one indeed does not have to worry about

the nomenclatural consequences caused by lumping or splitting any well-supported

clade can be named without affecting the name of more or less inclusive clades

Unfortunately unflagged hierarchies have very serious deficiences as communication

devices because they lack one aspect essential in language biological or otherwise -

they contain no intrinsic information about the relationships of the group in question to

others (eg Pfeil amp Crisp 2005 Stevens 2006a) Recent suggestions for using prefixes like

Apo- and Pan- to PhyloCode names will however allow limited information of this

kind to be conveyed but only as it pertains to individual branches and current

proposals do not even mandate that the prefixes be employed consistently In any

event such proposals simply prevent the potential tripling of the number of quite

different names used to describe different aspects of a phylogenetic tree over those

used to name monophyletic groups pure and simple In general where n is the number

of extant species in a group the number of clades in such a group = n-1 (Species will

also need names too for their names see Dayrat et al 2008) Importantly here too

consensus over the clade names commonly learned by students and used in herbaria is

needed otherwise communication will be impeded the names themselves will provide

no guidelines as to which should be chosen The situation is of course more complicated

than this Terminations that convey ideas of rank in a phylogenetic classification can also

be used in phylocode names - however there they will carry no implications of rank

How they will be used is another matter of course

Of course there are other philosophies of classification and some still prefer

evolutionary classifications There classificatory principles differ substantially from those

followed here eg the recognition of paraphyletic taxa may be permitted however

detailed reasons for prefering the taxa that are recognised are rarely given although

nature and natural groups are often mentioned (cf Stuessy amp Koumlnig 2008) For

summaries of commonly used systems see Brummitt (1992) and Mabberley (2008)

new evolutionary systems appear every year or so However even those who allow or

promote the recognition of paraphyletic groups (eg Grant 2003 Thorne 2007

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

13

Heywood et al 2007) may find it of interest to examine a system recognizing only

monophyletic groups it provides a rather different understanding of evolution

Evolutionary classifications in general try and combine phylogeny and

morphological gaps although that is no easy thing to do - it is akin to combining chalk

and cheese (for an attempt to make this impossible task seem to be more objective see

Stuessy amp Koumlnig 2008) However there is a principle from evolutionary classification

that is relevant and I think quite useful here The size of the gap between two groups

tends to be inversely proportional to the sizes of the groups involved (Davis amp Heywood

1963) One can imagine a situation in which a large group is formally divided even

although the distinguishing characters of the two are weak whereas a smaller group

similarly divisable will be left intact

To summarize If hypotheses of phylogeny remain stable we should be able to base

a stable classification on that phylogeny and then get on with our work that is testing

the phylogenies we have elucidating phylogenies in areas where relationships are

unclear studying the evolution of morphology describing species etc In this context

the spread of the Angiosperm Phylogeny Group system (see below) and its widespread

utilisation in technical literature also floras (eg van der Meijden 2005) textbooks (eg

Simpson 2006 Judd et al 2007 [third edition]) dictionaries (Mabberley 2008) more

popular literature (eg Souza amp Lorenzi 2005 Spears 2006) and as an outline for a new

herbarium sequence (Haston et al 2007) is gratifying The posibility that one might be

able to develop a stable phylogeny-based classification of families and in particular

orders represents a dramatic turn-around from the pessimistic attitude about such

higher-level groupings expressed by Davis and Heywood (1963) and Thorne (1976) the

latter even suggesting that we should bury forever the metaphor of the phylogenetic

tree as highly unrealistic (ibid p 56) Returning to Godfray and Knapps (2004) users

of classifications who want a stable informative and accessible classification that

enables easy identification - unfortunately they want cake with everything and cannot

get it - these pages attempt to satisfy as many of their needs as possible but phylogeny

and monophyly are the primary shapers of the APG classification

On this classification in particular

Here I very largely follow the Angiosperm Phylogeny Group classification (APG

2003) Any differences are not to be interpreted as differences in principle simply that

new phylogenies continue to be published and here I attempt to provide an overview

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

14

of current ideas of higher-level relationships of seed plants The Angiosperm Phylogeny

Group classification is based on well-supported relationships evident in the numerous

molecular studies that began to appear in the late 1980s (see APG 1999 for the

principles underlying the classification) but recent phylogenetic work does not

contradict the major outlines of the trees used by APG II (2003) or even those in APG

I (APG 1999) The APG classification has for the most part been conservative and I

am similarly conservative here For the names and authorities I follow APG III (2009)

although the names of the authors may not always be correct these being in something

of a state of flux for the superordinal names used see Chase and Reveal (2009)

although classification at this level here is not exhaustive - only clades with several

orders have superordinal names

Some changes in our ideas of relationships and hence in the clades we talk about

are particularly likely in parts of Lamiales Caryophyllales and Malpighiales for example

although this will not affect the orders themselves Thus some changes to clade

circumscriptions are to be expected although I do not expect them to be substantial

but changes are neither a defect of cladistics nor a necessary consequence of the use of

molecular data Clades are hypotheses of relationships and as hypotheses they may be

overturned However work since APG I has not suggested other than minor changes

in the compositions of the orders even if the odd genus or even family is turning out to

be seriously misplaced - recent examples are Hydatellaceae (from monocots-Poales to

Nymphaeales Saarela et al 2007) Guamatelaceae (from Rosales-Rosaceae to

Crossosomatales Oh amp Potter 2006) and Perrottetia and Bhesa (from Celastrales-

Celastraceae to Huerteales and Malpighiales respectively Zhang amp Simmons 2006) The

main changes have been clarification of the relationhips of individual families or groups

of families that were of uncertain position eg of Chloranthaceae (Moore et al 2007)

Ceratophyllaceae (Jansen et al 2007) and unplaced asterid II families (Winkworth et al

2008a) As mentioned above phylogenies in the area of main interest in these pages are

overwhelmingly tree-like Although there is evidence for acquisition of host plant genes

by parasites (see below) even wider but inexplicable transfer of mitochochondrial

genes in plants like Amborella (Bergthorsson et al 2004 cf Goremykin et al 2009a)

and even transfers of nuclear genes (Vallenback et al 2008) these are the exception

rather than the rule

In cases where the Angiosperm Phylogeny Group II allowed alternatives as to the

limits of families - Papaveraceae in the broad sense or Papaveraceae plus

Pteridophyllaceae plus Fumariaceae Proteaceae in the broad sense or Proteaceae plus

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

15

Platanaceae - the choices made here follow common usage eg as in textbooks like

Judd et al (2007) and Simpson (2006) and particularly in the new edition of

Mabberleys The Plant Book (Mabberley 2008) This latter is itself an attempt to reflect a

consensus the result of taking the opinions of botanists at several meetings A largely

similar consensus is reflected in the APG III classification (APG 2009) where you will

see that alternative classifications have been dispensed with For many people the

existence of such alternative classifications will simply confuse so agreement over

which groupings to use when alternatives are permitted and then dispensing with the

alternatives that are not commonly used seems reasonable I am making the few

changes that are needed to bring the classification here in line with these others but

only gradually since there is not enough time in each day

As already mentioned higher-level relationships in general and the composition of

orders in particular have always presented something of a challenge to systematists

(eg Davis amp Heywood 1963) That being said the composition of some clades like

Apiales Crossosomatales and Pandanales is decidedly unexpected However it is

interesting to see that these higher level clades are generally accepted even in works

with different classificatory philosophies (eg Heywood et al 2007) For clades like

Malpighiales many of the family groupings within Asparagales etc attempts to find

distinctive characters have largely failed (but see Endress amp Matthews 2006a also

the Apomorphies page here) Interestingly as with families some groupings suggested

by molecular studies are supported by morphological andor chemical characters that

have long been known sometimes for over a hundred years the relationship between

Pittosporaceae and ApiaceaeAraliaceae are a case in point (Hegnauer 1969b and

references) As our knowledge of morphology and chemistry improves we can hope for

improvements in the characterisations of clades at all levels

In many cases the new family limits of the Angiosperm Phylogeny Group (see

APG 1999 2003) are not really controversial although changes from the limits

commonly accepted only a decade ago are sometimes quite dramatic (eg Wagenitz

1997) Thus the split of the old Saxifragaceae sl is necessitated by its extreme

polyphyly as also with Icacinaceaesl and Cornaceae sl However the limits and

relationships of such groups had long been considered to be unsatisfactory but until

recently there had been no real reason to prefer one proposed arrangement over

another It is generally accepted that the limits of Lamiaceae and Verbenaceae have to

be redrawn although there is as yet no compelling evidence that the redrawn taxa are

not sister taxa (there is no evidence that they are) But whatever their relationships the

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

16

content of the clades has changed considerably and incidentally they are now easier to

identify than before the decision to recognise two families is not difficult The same is

true for Salicaceae and Achariaceae (Malpighiales) two previously small families that

have received the bulk of the old Flacourtiaceae Clade and hence taxon limits remain

difficult around Theaceae (Ericalessee alsoSladeniaceae Pentaphylacaceae)

and Euphorbiaceae (Malpighiales see

also Phyllanthaceae Picrodendraceae Putranjivaceae etc) Here current groupings

may not represent quite such dramatic changes in our understanding of relationships

Indeed parts of the old Euphorbiaceae that were separated may yet go back together

particularly Phyllanthaceae and Picrodendraceae (see Wurdack et al 2004 Davis et al

2005) even if Theaceae should indeed be dismembered (Geuten et al 2004) However

given that there is no molecular evidence that currently warrants combining all the

segregates of Euphorbiaceae and even if some do come together the clades the

families represent suggest novel groupings not recognised in current classifications

Furthermore Rafflesiaceae appear to be embedded within Euphorbiaceae s str (Davis

et al 2007) so the family is divided Of course maintaining even a moderately narrowly-

circumscribed Euphorbiaceae would entail reducing the iconic Rafflesiaceae to

synonymy Relationships in core Caryophyllales especially

around Phytolaccaceae andNyctaginaceae and also Portulacaceae are incompletely

understood and refashioning of taxon limits will doubtless be needed as cladistic

relationships become apparent (eg see Nyffeler 2007 Ogburn 2007a b) Some

groupings in the old Icacinaceaeand Olacaceae also remain unclear

The discovery of the relationships of parasitic and aquatic groups have presented a

particular challenge to systematists Morphologically some of these plants are so highly

modified that interpretation of the plant body in conventional terms is difficult or even

impossible Thus parasitic groups (broad-leaved angiosperms only) such as Rafflesiaceae

are hard to place since both the vegetative body and the flowers are changed almost

beyond recognition (flowers of rafflesiaceae are a good example of gigantism - Davis et

al 2007) Furthermore plastid gene sequences may be difficult or impossible to obtain

the chloroplast DNA in particular being highly degraded and the rate of molecular

change in general may be high so the problem of long-branch attraction is serious (eg

Duff amp Nickrent 1997 Nickrent et al 1998 Caddick et al 2002a G Petersen et al

2006b) Echlorophyllous sapromycoheterotrophic taxa mostly known from monocots

present similar problems However progress is being made placements for

Apodanthaceae Rafflesiaceae Mitrastemonaceae Cytinaceae and Cynomoriaceae have

recently been suggested (Barkman et al 2004 2007 Davis amp Wurdack 2004 Nickrent et

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

17

al 2004 Davis et al 2007) it seems likely that Burmanniaceae sl are polyphyletic but

both parts are in Dioscoreales (Merckx et al 2006) and relationships within the largely

hemiparasitic Santalales are also gradually being clarified (Maleacutecot 2002) The inclusion

of parasitic taxa in general molecular analyses can cause conniptions (eg Nickrent et al

2004 Davis et al 2004 Chase et al 2006 G Petersen et al 2006b) and there can be

horizontal transmission of genes (eg Davis amp Wurdack 2005 Vitaceae to Rafflesiaceae

Barkman et al 2007 the mitochondrial atp1 gene commonly moves)

Similarly in water plants neither vessels in particular nor much xylem in general is

needed leaves are highly modified and water-mediated pollination if adopted may

well be associated with major changes in floral morphology Here too recent molecular

studies suggest that aquatic groups with hitherto problematic relationships may find

homes Thus Podostemaceae are close to Clusiaceae and Calophyllaceae and are sister

to Hypericaceae (Malpighiales Kita amp Kato 2001) Hydatellaceae which used to be in

Poales are part of Nymphaeales (Saarela et al 2007) and Hydrostachyaceae may be

close to Hydrangeaceae (Cornales Xiang et al 2002) or perhaps should be placed in

Lamiales note that in the first two cases in particular there are morphological and

chemical features that support such a move If Podostemaceae are indeed close to

Clusiaceae I look forward to seeing hypotheses to explain how the dramatic changes in

the vegetative body that have made Podostemaceae so problematic for generations of

systematists took place That conventional wisdom has trouble in understanding or

explaining how the morphologies of groups like Clusiaceae and Podostemaceae can be

related is largely a problem with conventional wisdom Ceratophyllaceae seem to be

finding a position as sister to eudicots (see Moore et al 2007) but their morphology is

so derived that there is no morphological evidence of which I am aware for this

relationship There are also similar although less extreme morphological problems with

many plants pollinated by wind

In such situations especially with some water plants mycoheterotrophs and

parasites a variety of characters may be affected and particularly under such

circumstances groups have tended to be formed using a few or even only a single

character that seems to provide evidence of relationships Add to this the tendency to

weight some characters particularly strongly on a priori grounds and the result was

either the recognition of conglomerate taxa such as Amentiferae which now appear to

be highly polyphyletic or the segregation of families like Plantaginaceae s str (now

much expanded) and Leitneriaceae (now in Simaroubaceae) - all of these are more or

less wind pollinated

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

18

Even although relationships of many parasitic and aquatic groups are being

clarified and at least some of the proposed phylogenetic positions seem to be well

supported it is indeed often still very difficult to understand how such groups have

evolved from their more morphologically conventional relatives However we have had

all sorts of preconceptions as to how characters should evolve Thus we used to

assume that features like highly scalariform vessel perforation plates or the complete

absence of vessels or a flower with an androecium that had many stamens a superior

ovary or separate petals were necessarily plesiomorphic or primitive and conversely

a simple perforation plate and the presence of vessels an androecium with few

stamens inferior ovary or petals that were connnate were almost necessarily

apomorphic or advanced (but cf eg Stebbins 1951) Such assumptions are incorrect

(eg Soltis et al 2005b) Carpels may become secondarily free carpels may fail to close

the seeds then developing outside the confines of the carpel as in some Aspagaraceae-

Nolinoideae Violaceae Berberidaceae Malvaceae-Sterculioideae etc) in Peliosanthes

teta perhaps the only species in Peliosanthes(Asparagaceae-Nolinoideae) the ovary

varies from superior to inferior (Jessop 1976 species limits here need close

investigation see also Kuzoff et al 2001 and Soltis amp Hufford 2002 Saxifragaceae

Apiales etc) many-seeded carpels can evolve from few-seeded carpels

(Razafimandimbison et al 2008) monoecy may be derived from dioecy (Schaefer amp

Renner 2010 and references) Classic studies such as those by Babcock (eg 1947)

on Crepis that assumed that evolution - in this case of the karyotype in particular - was

unidirectional have needed comprehensive re-evaluation (Enke amp Gemeinholzer 2008)

Most if not all characters have reversed andor evolved in parallel as is clear in the

discussion of ovary position here especially in Poales Saxifragales and Asparagales and

also in comments on such morphologically distinctive taxa (when compared with their

immediate relatives) such as Menyanthaceae and especially Pittosporaceae Parallel

evolution may occur even at the level of amino acid substitution as in the independent

acquisition of the phosphoenolpyruvate carboxylase (pepC) gene in C4 photosynthesis in

grasses (Christin et al 2007b see also Blaumlsing et al 2000)

Thinking of how such characters have been used in the past in classifications at the

generic level the results have been similar to those just discussed above for parasitic

plants and others Relying too much on animal pollination syndromes to mark generic

boundaries has all too often led to taxa that are highly unsatisfactory phylogenetically

(see eg Acanthaceae Bignoniaceae Campanulaceae Ericaceae Lamiaceae

Orchidaceae) and over-reliance on characters of fruit and seed (see eg Brassicaceae

and Apiaceae) has also led to unsatisfactory generic limits Again the more general

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

19

problem is the use of one or a very few characters that have been weighted a priori to

structure classifications (see also Garciacutea et al 2009) although it is perhaps not

surprising that such an approach sometimes results in the delineation of taxa that have

indeed turned out to be quite distinct (cf eg Burtt Astragalus versus Oxytropis)

Taxa recognized here are monophyletic yet many are polythetic at the

morphological level that is they lack unique features characterizing (defining) all and

only members of that family they can be recognised phenetically only by the unique

combinations of characters that they posess This is the result of evolution any

synapomorphy characterizing a taxon may be lost or modified beyond easy recognition

in some of its members or the synapomorphy may appear to be identical to a feature

that has evolved in parallel in a quite unrelated plant That plant groups are polythetic is

almost as much a feature of monophyletic taxa as the paraphyletic taxa common in

evolutionary classifications Some families now include substantial variation as

phenetically distinct derived groups are placed in their proper phylogenetic position -

examples are Ericaceae which include the erstwhile Empetraceae a wind-pollinated

group that is florally very different and the overwhelmingly large-flowered and animal-

pollinated Plantaginaceae which now include several derived small-flowered aquatic

and wind-pollinated groups that were previously placed in separate families

Thus some families as delimited here may not be easy to recognize However

remember that detecting relationships - use whatever characters you can even if they

are not obvious - and naming a plant - focus on easy-to-see characters that may not

reflect relationships - are quite different problems Taxa although natural may not be

readily recognizable indeed it was in exactly this context that Lamarck worked out the

basic principles of writing dichotomous keys in 1778 (see Scharf 2007 for keys and the

like) Of course Lamarcks idea of nature was very different from ours - he thought that

there was some kind of continuum of form on which living organisms were to be

situated with no real gaps anywhere - but this meant that his genera (for example)

might well not be sharply distinct from each other even if each were part of the real

continuum that was life Lamarcks separation of the establishment of relationships and

groupings in a system and identification of the plants in that system have been widely

accepted (but cf eg Godfray amp Knapp 2004 many of the contributors to Heywood et

al 2007) Perhaps the best way of identifying plants at the family level is by well-made

multiple access keys as in Watson and Dallwitz (1992a onwards family limits there may

differ substantially from those adopted here) Multiple access keys free users from the

constraints of dichotomous keys in which particular characters are needed at each step

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

20

of the identification process before there can be further progress Instead those

characters that happen to be evident on a specimen can be used in whatever order is

convenient when linked to illustrations glossaries etc their power is enormous (see

Dallwitz et al 2000 [2006] for the principles underlying their construction and use)

Nevertheless dichotomous keys such as those of Hutchinson (1973) and Franz Thonner

(Geesink et al 1981) have their uses Of course taxa coming out adjacent in keys may

well not be at all related

(When identifying large numbers of plants even more efficient than either style of

identification and certainly lots more fun is sight identification Unless you have a

photographic memory you have to build up your knowledge of comparative plant

morphology - on which the ability to make accurate identifications depends - by

repeated observation When faced with an unknown plant I always look for leaf teeth

and stipules distinctive hair types especially stellate and T-shaped hairs glands and

punctations of any sort the presence of latex or other exudate and check leaf insertion

smelling crushed leaves can also be helpful In this context nodal anatomy can usually

be checked using a razor and a hand lens or even simply carefully examining leaf scars

The short paragraphs added after most families may help in confirming familial

identifications)

For the record and for the little that it is worth there are 4 orders and 13 families

of gymnosperms characterised on these pages and together they include some 82

genera and 947 species For angiosperms comparable figures are 56 orders 445

families 13208 genera and 261750 species (of which monocots include 11 orders 89

families 2759 genera and 52760 species) Note however that higher mathematics

was never my strong point and anyway these are pretty meaningless figures even for

species which many (but not all) might concede smacked slightly more of reality than

other taxa estimates range as high as 422000 (Govaerts 2001) Furthermore numbers

of genera and species change daily Nevertheless as emphasized here families are

useful in teaching we as a community can ensure that their limits remain largely stable

and by concentrating on relatively few of them one can gain some familiarity with much

of the worlds flora For summaries of the sizes of orders and families and of the general

arrangement followed in these pages see also the Statistics page itself

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

21

A PHYLOGENETIC TREE OF SEED PLANTS (THE MAIN TREE)

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

22

The Main Tree above is a conservative summary of well supported relationships

between the major clades of extant seed plants Reading - interpreting and

understanding - such a phylogenetic tree is not necessarily easy so reading a tree may

require a bit of practice OHara (eg 1988 1992) provides an accessible introduction to

thinking and talking about history in the context of looking at phylogenetic trees More

recently Krell and Cranston (2004) Crisp and Cook (2005) and others have emphasized

how careful one must be when interpreting and talking about evolution in ladderized

trees in particular and phylogenetic trees in general Gregory (2008) provides a

particularly detailed discussion of this problem In ladderized trees the smaller (in terms

of numbers of terminals) sister taxon is consistently shown on the same side at every

node thus the trees tend to be pectinate like the teeth of a comb however the

horizontal axis that is as a result so evident has no polarity or particular direction The

use of the adjective basal when discussing phylogenetic trees is especially dangerous

(see also Wojciechowski et al 2004 D Soltis et al 2005b) When I use the term and the

context is not otherwise clear I am referring to the pectinations at the base of a

ladderised tree Remember that when talking about sister taxa one can never be basal

to the other or older or younger although members of one can be more derived - in the

sense of having more apomorphies - than the other (but even then one has to be

careful) The words primitive and advanced and lower and higher should

practically never be used especially when talking about taxa Amborellaceae and

Pinaceae are sister to all other flowering plants and Pinales and are basal to the crown

groups of all other flowering plantsPinales respectively but that does not mean they

are thereby primitive Pinaceae in particular have numerous apomorphies The word

plesiomorphic is far less loaded than primitive and can be used to talk about

individual characters

Most trees here have been more or less ladderized as is the one above The

ladderization may be imperfect for example you can see that on the Main Tree

asterids with ten orders follow rosids and relatives which have seventeen although

this is in part because of the currently unresolved nature of relationships between the

main clades of core eudicots (this is likely to change soon) The Main Tree could have

been drawn with Amborella Acorus or a host of other taxa at the far right without

offending any relationships phylogenetic trees are like mobiles the only fixed points

being the nodes Despite the problems just mentioned pectination interpreted

carefully has its value and it is not an arbitrary process As one reads the terminals of a

pectinate tree from left to right adjacent terminals will be separated by apomorphies

that will cumulate in an appropriate fashion When thinking of a book or a herbarium

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

23

sequence (see Haston et al 2007) this is of some value Nymphaeales and

Austrobaileyales are here adjacent on the tree but they could be separated by

hundreds of families in the sequence merely by rotating only the node from which

Nymphaeales and their sister group (all other angiosperms minus about 75 species)

arise If adjacent in a book or herbarium or as in these summary pages then it is

relatively easy to relate their characters both apomorphic and plesiomorphic but if

separated by hundreds of pages or two floors in a large building then it is less easy to

get anything from the sequence (Since all orders in the pages of the main part of this

site are preceded by the apomorphies of all nodes immediately below them in the seed

plant phylogeny and because of the linkages that have been built in to the site this

problem is much less serious there there is no sequence) So the particular sequence of

all seed plants that is based on a phylogenetic classification but is to be used in herbaria

or any other place where taxa have to be arranged linearly as in this part of APweb

will be one that maximizes the number of taxa that are both successive branches of the

tree and placed successively in the sequence Since specimens are generally filed under

families the outline of a new family sequence for arranging herbaria and books can now

be suggested (Haston et al 2007) and such a standardized sequence will further help

teaching and learning about plants

ON THINKING ABOUT APOMORPHIES

Identifying apomorphies is important because understanding the evolution of

morphology in the broad sense - ie including anatomy chemistry etc - in the context

of diversification is one of our major goals For identifying apomorphies several

preconditions must be met One needs to have an accurate well supported phylogeny

one has to have examined the right taxa both from the point of view of morphology and

molecules one has to have coded the characters right (ie delimited states

appropriately) and one has to use an appropriate model of evolution when fitting the

variation to the tree (see eg Omland 1999 Stevens 2006b) If the distributions of

apomorphies below are compared with those in other studies (eg Turgeon et al 2001

Bremer et al 2001 Endress 2001 Albach et al 2001a Judd amp Olmstead 2004 D Soltis

et al 2005b Zhang et al 2006 Judd et al 2007) the reader may find a number of

differences Although I have integrated such studies as far as is possible there are five

reasons that link to the preconditions just mentioned why there may be differences

where features are placed on trees

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

24

1 I may not have found all the information about a particular character there may be

disagreement over its interpretation or I have added information

2 The sampling of nearly all molecular studies is very incomplete (see Salisbury amp Kim

2001 for problems caused by sampling) indeed it is commonplace to decry the

incompleteness of molecular sampling But not only is the sampling in molecular studies

often less than we might wish that of the morphological and chemical characters whose

evolution we are interested in understanding is also often very poor So for many

anatomical chemical and embryological characters that are confidently said to

characterise families and other groups we all too often have no idea if those characters

are applicable to the whole clade or just to a subgroup within that clade However as

our sampling improves we can locate changes on the tree more precisely if in

unexpected positions Thus is is sometimes suggested that Ericaceae have ellagic acid

(Soltis et al 2005b) in fact the little evidence we have indicates that only a small clade

of some 80 species has ellagic acid and the rest of the family - itself alone about one

third of all Ericales - does not Similarly Ericaceae are noted for their

ectendomycorrhizal associations but Enkianthus sister to all other Ericaceae

apparently lacks these (Abe 2005) Thus these mycorrhizae and their associated

distinctive hair roots are unlikely to be apomorphies of Ericaceae - however it appears

that other families near Ericaceae may also have ectendomycorrhizae (see Asai 1934)

3 I am fitting characters to a very conservative tree with many polytomies although the

nodes that are utilised are for the most part strongly supported polytomies make the

optimisation of characters that is the assigment of character state change to a

particular node on the tree notably difficult (eg Madison amp Madison 2002) In nearly

all studies of the evolution of characters distributions of characters are optimised on a

more or less fully resolved tree and the construction of supertrees may yield yet more

detailed hypotheses of relationships (for literature on supertrees see Cotton amp

Wilkinson 2007 2008) Of course some nodes on such fully resolved trees andor

supertrees may have little support and optimisations of characters on such trees may

carry correspondingly little conviction Even parts of some of the trees used here have

poor support eg relationships within aquatic Alismatales etc although I indicate

when this is the case

4 Exactly how one goes about optimising a character on a tree is critically important

Even using simple parsimony optimisations (ACCTRAN or DELTRAN ACCelerated

TRANsitions or DELayed TRANsitions) the position of synapomorphies on trees - and

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

25

hence our ideas of evolution - may differ and this is still more true if one uses maximum

likelihood or Bayesian methods and makes apparently reasonable suggestions about

weighting gains over losses (eg Donoghue amp Ackerley 1996 Cunningham et al 1998

Omland 1997 1999 Ree amp Donoghue 1999 Polly 2001 Webster amp Purvis 2001

Ronquist 2004 Crisp amp Cook 2005) Sannier et al (2007) show how in in Arecaceae that

where on a phylogenetic tree one might peg changes in microsporogenesis will depend

on the methods one uses to do this and Pedersen et al (2007) discuss the sometimes

very substantial effect of node support on the posterior probabilities of ancestral

character states Here I use parsimony optimization not always as explicit as it might be

but I have often indicated where there are particularly important uncertainties as to the

positions of particular character changes on the tree

5 Finally although I have paid quite a lot of attention to the delimitation of the

character states that make up all the characterizations I have not spent enough time on

this critical operation If we are interested in understanding evolution then fitting the

basic variation - not character states - to a tree in principle allows greater flexibility in

understanding morphology in the context of local phylogenies (see also Stevens 2000

Endress 2005c) However many character states used here are delimited globally that

is they are circumscribed in the context of the variation shown by individual characters

across all angiosperms andor in the context of classic ideas of character evolution

Character states often have arbitrary limits and serve best to communicate

information whether they are in fact suitable for either phylogenetic analysis or

understanding evolution are separate issues Studies have rather unsurprisingly

perhaps but importantly shown that dividing the one character into different sets of

states may yield differing ideas of evolution of that character (eg Lamb Frye amp Kron

2003 Hibbett 2004) When looking at trees on which character states are optimised

one should bear in mind the problems surrounding the delimitation of states (eg

Stevens 2000 2006b) and the danger of using pollen or other types - constructs

based on many characters that vary independently but which effectively get lost in

these types thus Blackmore et al (2009) decomposed the pollen types in Asteraceae

into 52 characters

I use Remanes three main criteria of homology or better similarity when

determining the basic similarity of structures on different organisms (see Remane 1952)

These criteria are special properties position and intermediates Special properties

include anatomical or chemical characters gene expression data etc Position refers

to the position of an organ with respect to landmarks on the plant Although plants are

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

26

plants and landmarks are not as fixed as one might like trying to work out the relative

positions of parts is a good way of understanding morphology so long as one does not

try to out-do Procrustes Intermediates include intermediates found on the same

plant or on different plants Intermediates may observed during development or be

adult structures Thus one may find changes during development which suggest what

the nature of a particular structure is and very different stuctures on different plants

may turn out to be similar early in development Or one can simply compare different

taxa and see that structures that appear to be very different when comparing two

different taxa can be linked morphologically by looking at other taxa

Nevertheless the use of these criteria may not yield an unambiguous answer as to

what a structure is even given a solid phylogeny and an improved understanding of

development (see Jaramillo amp Kramer 2007 for a useful discussion) As Endress (2005c)

observed a number of features - position function development shape anatomy

histology gene activity and relationships to other taxa that clearly have petals - can be

used to distinguish a petal (for example) from other floral structures if a petal does not

have one of these features is it thereby not a petal Thus Maturen et al (2005) recently

found that floral organ diversity genes (B and C) were expressed in the large white

inflorescence bracts of Cornus (see also Costa et al 2005) Peney et al (2005) noted that

not all monosulcate pollen grains in monocots have the same developmental pathway

and that as a result such pollen might not have the same ancestral state Reeves and

Olmstead (2003) suggested that the genetic mechanisms causing monosymmetry in

Lamiales and Solanales were different and Serna and Martin (2006) described similar

problems with the development of hairs in Arabidopsis when compared with that of

hairs in Antirrhinum and Solanaceae Indeed as one perhaps might expect delimitation

of states and characters does not necessarily become easier with increasing knowledge

of development etc Thus Buzgo et al (2004) Matthews and Endress (2005) and others

have shown how hard it can be to distinguish between eg prophylls and other floral

structures as their behaviour is studied during the course of the development of the

flower Similarly at what concentration is a particular secondary metabolite deemed to

be present (Waterman (2007)

The validity of the approach used here that of fitting morphological variation to a

largely molecular-based tree may be questioned However I think it rather unlikely that

well-supported molecular branches will be overturned by morphological data Indeed

analyses of morphological data alone do provide support for many of the clades evident

in molecule-only analyses and analyses with morphological and molecular data

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

27

together may lead to increased support for clades (eg Hufford 1992 Nandi et al 1998

[but adding morphological data reduces support for a number of critical clades too]

Doyle amp Endress 2000 however in none of these papers is the use of morphology

without ambiguity) It is unfortunately clear that the use of morphology alone may not

suggest problems in the phylogenetic placement of taxa that later turn out to have been

wrongly included (eg Zhang et al 1992) Thus I have been wary of putting much weight

on clades that have only morphological support but note that I have treated molecule-

based clades with low bootstrap or jacknife support values (esp below 70) or low

posterior probabilies (below 095) likewise Although I may have been mistaken in

placing so much emphasis on molecular data in terms of providing the basic

phylogenetic framework for angiosperms morphological and molecular data are only

very rarely in irreconcilably strong conflict There are indeed a few places where the

conflict seems extreme These include the relative positions of the Monimiaceae and

Hernandiaceae (Laurales) the position of Hanguanaceae (Commelinales [as here] or

Zingiberales) and of Triplostegia (is it in Dipsacaceae or Valerianaceae - see

Dipsacales) Fossils are unlikely to affect the topologies of the trees presented here but

see below for their importance in understanding morphological evolution in general and

the evolution of angiosperms in particular Nevertheless some largely reject the idea

that trees based on molecular data alone can recover phylogenetic relationships

especially when branching points are old and prefer to used trees based on analysis of

morphological data including those taken from fossils (Hilton amp Bateman 2006 Farjon

2007)

All in all however the extent of the congruence between morphological and

molecular data is impressive and heartening and many clades can be characterised

morphologically It seemed in 1998 that there were no unambiguous morphological

synapomorphies for angiosperm orders (K Bremer 2000) and this is still true if by

unambiguous is meant non-homoplasious However many orders and other clades

have synapomorphies even if these may be indistinguishable at least at the current

level of morphological and developmental knowledge from parallel occurences

elsewhere As our morphological knowledge increases so too does the number of

apomorphies

Endress and Matthews (2006a) emphasize the importance of tendencies and

developmental constraints when thinking about characters of clades within the rosids -

many characters or character combinations occur in a rather sporadic fashion within

the clade and apparently notably less frequently outside it There are a number of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

28

examples of tendencies here and in the individual order pages such as the features

enclosed by parentheses in groups above the level of family eg see (cuticular waxes

as aggregated rodlets) for the commelinids and the discussion of the distribution of

polyandry that is flowers with numerous stamens in the asterid I + II groups etc

Tendencies may involve more than single characters When thinking of the

associations of features like integument thickness and vascularization of the integument

(Endress amp Matthews 2006a) one can imagine fairly simple developmental

preconditions being operative As Endress and Matthews (2006a) note it is difficult to

imagine vascular tissue developing in an integument only two cells thick Changes here

would be loosely correlated if morphologically linked However the strongly correlated

changes noted by Givnish et al (2005) are ecologically linked but are presumably

morphologicallydevelopmentally independent When there seem to be characters

evolving more or less together tests can be carried out to see if the changes are

concentrated on certain branches of the tree (eg Maddison 1990 Sanderson 1991

Maddison amp Maddison 2000) Such tests have rarely been carried out at higher levels in

angiosperms

One of the most striking examples of a tendency is the distribution of N-fixation

restricted as it is to a monophyletic group of four clades although it has arisen seven (or

perhaps several more) times independently within the N-fixing clade and several

members of two quite different kinds of bacteria are involved (eg Clawson et al 2004

Elliott et al 2007 Sprent amp James 2007) Other examples of work that bears on the issue

of tendencies include the findings that flowers of polysymmetrical Arabidopsis have

genes like TCP1 that are expressed asymmetrically during early development and TCP1

is a probable orthologue of the well-known CYC gene of Antirrhinum that is involved in

the development of monosymmetric slowers there (Cubas et al 2001 Costa et al 2005

etc) Parallelism might build on this underlying morphologically cryptic monosymmetry

even if details of the genetic mechanisms causing the monosymmetry evident in

particular groups may be different (Reeves amp Olmstead 2003 Cubas 2004 see above)

Similarly there have been several recent suggestions that the capability to synthesise a

particular metabolite may be switched off but not lost and so can sometimes be

reacquired (eg Wink amp Witte 1983 Wink 2003 Liscombe et al 2005 Larsson 2007

Waterman 2007) Hence perhaps the rather spotty distribution of many secondary

metabolites like ellagic acid the indole alkaloid camptothecin iridoids etc when

considered in the context of phylogenies As a non-botanical example - but a rather nice

one - Salwini-Plawen and Mayr (1961) suggested some time ago that there has been

considerable parallelism (40-65 or more independent origins) in the evolution of eyes in

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

29

metazoans However Pax 6 seems to be a master control gene that is involved in eye

formation perhaps throughout all bilateralians (eg Gehring amp Ikeo 1999 Erwin amp

Davidson 2002) which changes the nature of the problem Note however that in

general our invocation of evolutionary tendencies to explain the patterns of

distributions of characters is really an admission that we do not understand those

patterns

As we find out more about variation we will probably find fewer and fewer features

constant throughout a group Most unqualified statements of presence and absence

should properly be qualified as usually present or usually absent if one is thinking of

the characterisations on the order pages as encompassing the total variation within a

clade Thus Pistia alone among monocots as so far known has sieve tube plastids with

starch grains not protein crystals However this does not affect the fact that sieve tube

plastids with cuneate protein crystals and lacking starch grains are an apomorphy for

monocots Furthermore fossils in a number of cases suggest character combinations

unknown in extant taxa as may be seen in the discussions of Fagaceae Platanaceae

Iteaceae Calycanthaceae etc Confusing the issue there may be questions as to where

exactly on the tree a particular fossil is to be placed (eg see Nymphaeaceae

Calycanthaceae Archaefructus etc)

To summarize given our current understandings of both phylogenies and

characters evolution of some characters in which we are interested seems very labile

(see eg D Soltis et al 2005b Endress and Matthews 2006a Stevens 2006b Ekman et

al 2008 for a good recent study) and I have been cautious when talking about character

evolution Much effort must continue to be spent in summarizing characters of well-

established clades at all levels providing features by which they may be recognized and

signaling synapomorphies Remember that (1) the basic morphological anatomical and

chemical knowledge of many critical taxa is woefully incomplete (2) different

assumptions about character evolution may greatly affect the position of

synapomorphies on trees (3) in many cases relationships within and between many

groups are too uncertain at present to worry very much about synapomorphies and

(4) we must be clear about what we do and do not not know As mentioned above I

have indicated in a number of places where there are particularly important

uncertainties as to where characters should be placed on the tree Nevertheless it is a

relatively easy matter to update notes such as these and it can be a simple matter to

incorporate new data on characters that have never before been considered in the

context of a tree Much basic - and unfortunately perhaps unfashionable - work must be

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

30

carried to clarify the distribution of morphological anatomical and chemical characters

for examples of what can to be done see the work of P K Endress and collaborators

(floral morphology and development) and S R Jensen and collaborators (iridoids) But

acquiring information about nodal anatomy cell and tissue distribution nodal anatomy

and the like is not difficult at all and should be generally encouraged

But all this is merely a necesary prelude to the understanding of evolution For this

one needs to know a lot more including dating the phylogeny understanding the

function(s) of characters factoring in relevant aspects of the palaeoenvironment in

which an apomorphy first appeared etc

SUMMARY OF THE SYSTEM

Below is a formalised summary of the relationships within orders of the families of seed

plants There are a few families that are not recognised even as options in APG II and

vice versa as well as a few extra orders The families to be recognised in the most

recent edition of Mabberleys The Plant Book (Mabberley 2008) are also largely

consistent with those below But all differences are trivial and will - I hope - eventually

disappear

Square brackets - [] - enclose clades the plus sign - + - designates sister taxa a

comma - - denotes part of a polytomy and quotation marks - - denotes a

paraphyletic group

SEED PLANTS

GYMNOSPERMS

Cycadales

Cycadaceae + Zamiaceae

Ginkgoales

Ginkgoaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

31

Pinales

Pinaceae [[Araucariaceae [Podocarpaceae + Phyllocladaceae]] [Sciadopityaceae

[Taxaceae + Cupressaceae]]]

Gnetales

Ephedraceae [Gnetaceae + Welwitschiaceae]

ANGIOSPERMSFLOWERING PLANTS

Amborellales

Amborellaceae

Nymphaeales

Hydatellaceae [Cabombaceae + Nymphaeaceae]

Austrobaileyales

Austrobaileyaceae [Schisandraceae + Trimeniaceae]

Chloranthales

Chloranthaceae

MAGNOLIIDS

Magnoliales

Myristicaceae [Magnoliaceae [[Himantandraceae + Degeneriaceae] [Eupomatiaceae +

Annonaceae]]]

Laurales

Calycanthaceae [[Siparunaceae [Gomortegaceae + Atherospermataceae]] [Monimiaceae

[Hernandiaceae + Lauraceae]]]

Canellales

Canellaceae + Winteraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

32

Piperales

[Hydnoraceae + Aristolochiaceae] [Piperaceae + Saururaceae]

MONOCOTS

Acorales

Acoraceae

Alismatales

Araceae [Tofieldiaceae [[Alismataceae [Hydrocharitaceae + Butomaceae]]

[Scheuchzeriaceae [Aponogetonaceae [Juncaginaceae [Maundiaceae [[Posidoniaceae

[Ruppiaceae + Cymodoceaceae]] [Zosteraceae + Potamogetonaceae]]]]]]]

Petrosaviales

Petrosaviaceae

Dioscoreales

Nartheciaceae [[Taccaceae + Thismiaceae] [Burmanniaceae + Dioscoreaceae]]

Pandanales

Velloziaceae Triuridaceae Stemonaceae [Pandanaceae + Cyclanthaceae]

Liliales

Corsiaceae [Campynemataceae [Petermanniaceae [Colchicaceae + Alstroemeriaceae]]

Melanthiaceae [[Philesiaceae + Rhipogonaceae] [Smilacaceae + Liliaceae]]]

Asparagales

Orchidaceae [[Boryaceae [Blandfordiaceae [Lanariaceae [Asteliaceae + Hypoxidaceae]]]]

[[Ixioliriaceae + Tecophilaeaceae] [Doryanthaceae [Iridaceae [Xeronemataceae

[Xanthorrhoeaceae [Alliaceae + Asparagaceae]]]]]]]

COMMELINIDS

Unplaced

Dasypogonaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

33

Arecales

Arecaceae

Poales

[Typhaceae + Bromeliaceae] [Rapateaceae [[Thurniaceae [Juncaceae + Cyperaceae]]

[[Mayacaceae [Eriocaulaceae + Xyridaceae]] [Flagellariaceae [Anarthriaceae

[Centrolepidaceae + Restionaceae]]] [Joinvilleaceae [Ecdeiocoleaceae + Poaceae]]]]]]]

Commelinales

[Commelinaceae + Hanguanaceae] [Philydraceae [Haemodoraceae + Pontederiaceae]]

Zingiberales

Musaceae [Strelitziaceae + Lowiaceae] Heliconiaceae [[Cannaceae + Marantaceae]

[Costaceae + Zingiberaceae]]

Ceratophyllales

Ceratophyllaceae

EUDICOTS

Ranunculales

Eupteleaceae [Papaveraceae [[[Lardizabalaceae + Circaeasteraceae] [Menispermaceae

[Berberidaceae + Ranunculaceae]]]]

Sabiales

Sabiaceae

Proteales

Nelumbonaceae [Platanaceae + Proteaceae]

Trochodendrales

Trochodendraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

34

Buxales

Haptanthaceae [Buxaceae + Didymelaceae]

CORE EUDICOTS

Gunnerales

Gunneraceae + Myrothamnaceae

Dilleniales

Dilleniaceae

Saxifragales

Peridiscaceae [[Paeoniaceae [Altingiaceae [Hamamelidaceae [Cercidiphyllaceae +

Daphniphyllaceae]]]] [[Crassulaceae [Aphanopetalaceae [Tetracarpaeaceae

[Penthoraceae + Haloragaceae]]]] [Iteaceae [Grossulariaceae + Saxifragaceae]]]]

Cynomoriaceae unplaced

Vitales

Vitaceae

ROSIDS

FABIDROSID I

Zygophyllales

Krameriaceae + Zygophyllaceae

Celastrales

Lepidobotryaceae + Celastraceae

Oxalidales

Huaceae [[Connaraceae + Oxalidaceae] [Cunoniaceae [Elaeocarpaceae [Brunelliaceae +

Cephalotaceae]]]]

Malpighiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

35

[Achariaceae [Goupiaceae [Violaceae + Passifloraceae] [Lacistemataceae Salicaceae]]

[Lophopyxidaceae Putranjivaceae] [Ctenolophonaceae [Erythroxylaceae +

Rhizophoraceae]] Linaceae Ixonanthaceae Humiriaceae Irvingiaceae

Centroplacaceae Pandaceae [[Bonnetiaceae + Clusiaceae] [Calophyllaceae

[Hypericaceae + Podostemaceae]]] [Malpighiaceae + Elatinaceae] Ochnaceae

[Peraceae [Rafflesiaceae + Euphorbiaceae]] [Phyllanthaceae + Picrodendraceae]

[Balanopaceae [[Trigoniaceae + Dichapetalaceae] [Chrysobalanaceae +

Euphroniaceae]]] Caryocaraceae

N-FIXING CLADE

Fabales

Quillajaceae [Fabaceae [Polygalaceae + Surianaceae]]

Rosales

Rosaceae [[Barbeyaceae [Elaeagnaceae Dirachmaceae Rhamnaceae]] [Ulmaceae

[Cannabaceae [Moraceae + Urticaceae]]]]

Cucurbitales

Anisophylleaceae [[Corynocarpaceae + Coriariaceae] [Cucurbitaceae [Tetramelaceae

[Datiscaceae + Begoniaceae]]]] Apodanthaceae

Fagales

Nothofagaceae [Fagaceae [[Myricaceae + Juglandaceae] [Casuarinaceae

[Ticodendraceae + Betulaceae]]]]

MALVIDROSID II

pgtGeraniales

Geraniaceae [[Melianthaceae + Francoaceae] [Vivianaceae + Ledocarpaceae]]

Myrtales

Combretaceae [[Onagraceae + Lythraceae] [[Vochysiaceae + Myrtaceae]

[Melastomataceae [Crypteroniaceae [Alzateaceae + Penaeaceae]]]]]

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

36

Crossosomatales

[Staphyleaceae [Guamatelaceae [Crossosomataceae + Stachyuraceae]]] [Aphloiaceae

[Geissolomataceae + Strasburgeriaceae]]

Picramniales

Picramniaceae

Sapindales

Biebersteiniaceae Nitrariaceae [Kirkiaceae [Anacardiaceae + Burseraceae]]

[Sapindaceae [Simaroubaceae Rutaceae Meliaceae]]

Huerteales

Gerradinaceae [Dipentodontaceae + Tapisciaceae]

Malvales

Neuradaceae [Thymelaeaceae [Sphaerosepalaceae Bixaceae [Cistaceae

[Sarcolaenaceae + Dipterocarpaceae]] [Cytinaceae + Muntingiaceae] Malvaceae]]

Brassicales

[Akaniaceae + Tropaeolaceae] [[Moringaceae + Caricaceae] [Setchellanthaceae

[Limnanthaceae [[Koeberliniaceae [Bataceae + Salvadoraceae]] [Emblingiaceae

[Pentadiplandraceae [Gyrostemonaceae Resedaceae Stixis and relatives] Tovariaceae

[Capparaceae [Cleomaceae + Brassicaceae]]]]]]]]

Berberidopsidales

Aextoxicaceae + Berberidopsidaceae

Santalales

Erythropalaceae Olacaceae [[Loranthaceae [Misodendraceae + Schoepfiaceae]]

[Opiliaceae + Santalaceae]] Balanophoraceae unplaced

Caryophyllales

[[Droseraceae [Nepenthaceae [Drosophyllaceae [Ancistrocladaceae +

Dioncophylleaceae]]]]] [[Frankeniaceae + Tamaricaceae] [Polygonaceae +

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

37

Plumbaginaceae]]] [Rhabdodendraceae [Simmondsiaceae [[Asteropeiaceae +

Physenaceae] [[Caryophyllaceae [Achatocarpaceae + Amaranthaceae]]

[Stegnospermataceae [Limeaceae [[Lophocarpaceae [Barbeuiaceae [Aizoaceae

[Sarcobataceae Phytolaccaceae Nyctaginaceae]]]] [Molluginaceae [Halophytaceae

Montiaceae Basellaceae Didiereaceae [Talinaceae [Portulacaceae [Anacampseros etc +

Cactaceae]]]]]]]]]]]]]

ASTERIDS

Cornales

[Cornaceae + Nyssaceae] [Hydrangeaceae + Loasaceae] Hydrostachyaceae [Curtisiaceae

+ Grubbiaceae]

Ericales

[Balsaminaceae [Marcgraviaceae + Tetrameristaceae]] [[Polemoniaceae +

Fouquieraceae] Lecythidaceae [[Sladeniaceae + Pentaphylacaceae] [Sapotaceae

[Ebenaceae [Maesaceae [Theophrastaceae [Myrsinaceae + Primulaceae]]]]]

[Mitrastemonaceae Theaceae [Symplocaceae [Styracaceae + Diapensiaceae]]

[[Sarraceniaceae [Roridulaceae + Actinidiaceae] [Clethraceae [Cyrillaceae +

Ericaceae]]]]]]

LAMIIDASTERID I

Unplaced

Metteniusaceae Oncothecaceae Icacinaceae and some unplaced ex-Icacinaceae

Garryales

Garryaceae + Eucommiaceae

Unplaced

Boraginaceae Vahliaceae

Gentianales

Rubiaceae [Gentianaceae [Loganiaceae [Gelsemiaceae + Apocynaceae]]]

Lamiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

38

Plocospermataceae [Oleaceae [Tetrachondraceae [Calceolariaceae

[Gesneriaceae Peltanthera [Pedaliaceae Carlemanniaceae Martyniaceae Stilbaceae

Plantaginaceae Linderniaceae Bignoniaceae Verbenaceae Lamiaceae Paulowniaceae

Schlegeliaceae Thomandersiaceae Phrymaceae Rehmannia Acanthaceae

Scrophulariaceae Orobanchaceae Byblidaceae Lentibulariaceae]]]]]

Solanales

[Montiniaceae [Sphenocleaceae + Hydroleaceae]] [Convolvulaceae + Solanaceae]

CAMPANULIDASTERID II

Aquifoliales

[[Cardiopteridaceae + Stemonuraceae] [Aquifoliaceae [Helwingiaceae +

Phyllonomaceae]]

Asterales

[Rousseaceae + Campanulaceae] Pentaphragmataceae [[Alseuosmiaceae [Phellinaceae

+ Argophyllaceae]] Stylidiaceae [Menyanthaceae [Goodeniaceae [Calyceraceae +

Asteraceae]]]]

Escalloniales

Escalloniaceae

Bruniales

[Bruniaceae + Columelliaceae]

Apiales

Pennantiaceae [Torricelliaceae [Griseliniaceae [Pittosporaceae [Araliaceae

[Myodocarpaceae + Apiaceae]]]]]

Paracryphiales

Paracryphiaceae

Dipsacales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

39

Adoxaceae [Diervillaceae [Caprifoliaceae [Linnaeaceae [Morinaceae [Dipsacaceae +

Valerianaceae]]]]]

LINKS TO ORDERS AND FAMILIES

[Back to Top]

MAIN GROUPINGS

Asterids commelinids core eudicots asterid 1 asterid

2 eudicots gymnosperms Magnoliophyta monocots N-fixing claderosids seed plants

ALPHABETICAL LISTING OF ALL ORDINAL NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | L | M | N | O | P | Q | R | S | T | U | V | W | X | Z

Abietales Acanthales Acerales Acorales Actinidiales Actinostrobales Adoxales Aescul

ales Agavales Aizoales AkanialesAlismatales Alliales Alseuosmiales Alstroemeriales

Altingiales Amaranthales Amaryllidales Amborellales AmbrosialesAmmiales Amomal

es Ancistrocladales Anisophylleales Annonales Anthobolales Apiales Apocynales Apo

nogetonalesAquifoliales Arales Araliales Aralidiales Araucariales Arecales Aristolochi

ales Athrotaxidales Asarales AsclepiadalesAsparagales Asphodelales Asparagales Ast

eliales Atriplicales Aucubales Austrobaileyales Avenales

Balanitales Balanopales Balanophorales Balsaminales Barbeyales Barclayales Batales

Begoniales BerberidalesBerberidopsidales Betulales Biebersteiniales Bignoniales Bix

ales Boraginales Brassicales Brexiales Bromeliales BrunialesBrunoniales Burmannial

es Burserales Butomales Buxales Byblidales

Cactales Callitrichales Calycanthales Calycerales Campanulales Campynematales Can

ellales Cannales CapparalesCaprifoliales Cardiopteridales Carduales Caricales Carle

manniales Caryophyllales Cassiales Casuarinales CelastralesCentrolepidales Cephalot

ales Cephalotaxales Ceratophyllales Cercidiphyllales Chenopodiales Chironiales Chlor

anthalesChrysobalanales Cinchonales Circaeasterales Cistales Citrales Cocosales Col

chicales Columelliales CombretalesCommelinales Connarales Convolvulales Coriarial

es Cornales Corylales Corynocarpales Crassulales CrossosomatalesCucurbitales Cunn

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

40

inghamiales Cunoniales Cupressales Cycadales Cyclanthales Cymodoceales Cynarales

CynomorialesCyperales Cyrillales Cytinales

Daphnales Daphniphyllales Dasypogonales Datiscales Degeneriales Desfontainiales D

iapensiales Didymelales DillenialesDioales Dioncophyllales Dioscoreales Diospyrales

Dipentodontales Dipsacales Droserales

Ebenales Echiales Elaeagnales Elaeocarpales Elatinales Elodeales Emmotales Empetr

ales Ephedrales Ericales EriocaulalesErythropalaless Escalloniales Eucommiales Euph

orbiales Eupomatiales Eupteleales Euryalaless

Fabales Fagales Falcatifoliales Ficales Flacourtiales Flagellariales Fouquieriales Franc

oales Frangulales

Galiales Garryales Geissolomatales Gentianales Geraniales Gesneriales Ginkgoales G

laucidiales Globulariales GnetalesGoodeniales Greyiales Griseliniales Grossulariales

Grubbiales Gunnerales Gyrocarpales Gyrostemonales

Haemodorales Haloragales Hamamelidales Hanguanales Heisteriales Helleborales He

lwingiales Himantandrales HippuridalesHomaliales Hortensiales Huales Huerteales

Hydatellales Hydnorales Hydrangeales Hydrastidales HydrocharitalesHydropeltidales

Hydrostachyales Hypericales Hypoxidales

Icacinales Illiciales Iridales Irvingales Iteales Ixiales

Jasminales Juglandales Julianiales Juncaginales Juncales

Lacistematales Lactoridales Lamiales Lardizabalales Laurales Lecythidales Ledocarpal

es Leitneriales LentibularialesLigustrales Liliales Limnanthales Linales Liriales Loasal

es Lobeliales Loganiales Lonicerales Loranthales LowialesLythrales

Magnoliales Malpighiales Malvales Marathrales Marcgraviales Mayacales Medusagy

nales Medusandrales MelanthialesMelastomatales Meliales Melianthales Meliosmal

es Menispermales Menyanthales Metteniusales MitrastemonalesMiyoshiales Moni

miales Moringales Musales Myricales Myristicales Myrothamnales Myrsinales Myrta

les

Najadales Nandinales Narcissales Nartheciales Nelumbonales Nepenthales Neuradal

es Nitrariales Nolanales NothofagalesNyctaginales Nymphaeales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

41

Ochnales Oenotherales Olacales Oleales Onagrales Oncothecales Opuntiales Orchid

ales Oxalidales

Saxifragales Pandales Pandanales Papaverales Paracryphiales Parasitaxales Paridales

Parnassiales PassifloralesPenaeales Pennantiales Pentaphragmatales Peridiscales P

etiveriales Petrosaviales Phellinales Philydrales PhyllanthalesPhysenales Phytolaccal

es Picramniales Pinales Pinguiculales Piperales Pittosporales Plantaginales Platanale

sPlumbaginales Poales Podocarpales Podophyllales Podostemales Polemoniales Pol

ygalales Polygonales PontederialesPortulacales Posidoniales Potamogetonales Primu

lales Proteales

Quercales Quillajales Quintiniales

Rafflesiales Ranunculales Rapateales Resedales Restionales Rhabdodendrales Rhamn

ales Rhinanthales RhizophoralesRhodorales Rhoipteleales Roridulales Rosales Rouss

eales Rubiales Ruppiales Rutales

Sabiales Salicales Salvadorales Samolales Samydales Sanguisorbales Santalales Sapin

dales Sapotales SarracenialesSaxegotheales Saxifragales Scheuchzeriales Sciadopitya

les Scleranthales Scrophulariales Scyphostegiales SedalesSilenales Simmondsiales S

milacales Solanales Sphenocleales Sphenostemonales Stangeriales Stellariales Stemo

nalesStilbales Stylidiales Styracales Surianales

Taccales Tamales Tamaricales Taxales Taxodiales Tecophilaeales Terebinthales Tern

stroemiales Theales TheligonalesThymelaeales Tiliales Tofieldiales Torricelliales Tov

ariales Tribelales Trilliales Trimeniales Triuridales TrochodendralesTropaeolales Turn

erales Typhales

Ulmales Urticales

Vacciniales Vahliales Vallisneriales Velloziales Veratrales Verbenales Viburnales Vinc

ales Violales Viscales VitalesVochysiales

Welwitschiales Winterales

Xanthorrhoeales Ximeniales Xyridales

Zamiales Zingiberales Zosterales Zygophyllales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

42

ALPHABETICAL LISTING OF ALL FAMILY NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | K | L | M | N | O | P | Q | R | S | T | U | V | W | X | Y

|Z

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lismataceae Alliaceae Allioniaceae Allophylaceae Aloaceae Alopecuraceae Alpiniacea

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ridaceae Apostasiaceae Aptandraceae AquifoliaceaeAquilariaceae Aquilegiaceae A

raceae Aragoaceae Araliaceae Aralidiaceaee Araucariaceae Arbutaceae Arceuthidace

aeArceuthobiaceae Arctostaphylaceae Arctotidaceae Ardisiaceae Arecaceae Argoph

yllaceae Arisaraceae AristolochiaceaeAristoteliaceae Arjonaceae Armeriaceae Artem

isiaceae Arthrotaxidaceae Artocarpaceae Arundinaceae ArundinellaceaeAsaraceae A

sclepiadaceae Ascyraceaee Asiraceae Asparagaceae Asperulaceae Asphodelaceae As

pidistraceaeeAsteliaceae Asteraceae Asteranthaceae Asteropeiaceae Astragalaceae

Astrocarpaceae AthanasiaceaeAtherospermataceae Atriplicaceae Atropaceae Aucuba

ceae Aurantiaceae Austrobaileyaceae Austrotaxaceae AvenaceaeAverrhoaceae Avet

raceae Avicenniaceae Azaleaceae Azimaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Balanitaceae Balanopaceae Balanophoraceae Balsameaceae Balsaminaceae Bambusa

ceae Barbaceniaceae BarbeuiaceaeBarbeyaceae Barclayaceae Barringtoniaceae Base

llaceae Bataceae Baueraceae Bauhiniaceae Baxteriaceae BegoniaceaeBehniaceae B

elangeraceae Belloniaceae Bembiciaceae Berberidaceae Berberidopsidaceae Berryac

eaee BersamaceaeBertyaceae Berzeliaceae Besleriaceae Betaceae Betulaceae Biebe

rsteiniaceae Bifariaceae Bignoniaceae BischofiaceaeBixaceae Blakeaceae Blandfordia

ceae Blattiaceae Blepharocaryaceae Blitaceae Blyxaceae Boerlagellaceae Bolivaracea

eBombacaceae Bonnetiaceae Bontiaceae Boopidaceae Boraginaceae Borassaceae B

oroniaceae BoryaceaeBotryodendraceae Bougainvilleaceae Boweniaceae Brassicacea

e Brachycaulaceae Bretschneideraceae BrexiaceaeBromeliaceae Brownlowiaceae Br

unelliaceae Bruniaceae Brunoniaceae Brunsvigiaceae Bryoniaceae BuchneraceaeBuci

daceae Buddlejaceae Buglossaceae Bulbocodiaceae Bumeliaceae Bupleuraceae Burc

hardiaceae BurmanniaceaeBurseraceae Butneriaceae Butomaceae Buxaceae Byblida

ceae Byttneriaceae

Cabombaceae Cacaoaceae Cactaceae Caesalpiniaceae Caladiaceae Calamaceae Calce

olariaceae CalectasiaceaeCalendulaceae Callaceae Callicomaceae Calligonaceae Calli

traceae Callitrichaceae Calochortaceae CalophyllaceaeCalthaceae Calycanthaceae Ca

lyceraceae Cambogiaceae Camelliaceae Campanulaceae CampynemataceaeCanacom

yricaceae Canellaceae Cannabaceae Cannaceae Canopodaceae Canotiaceae Cansjera

ceae CapparaceaeCaprariaceae Caprifoliaceae Cardiopteridaceae Carduaceae Carica

ceae Carissaceae Carlemanniaceae CarpinaceaeCarpodetaceae Cartonemataceae Ca

ryocaraceae Caryophyllaceae Casearieaceae Cassiaceae CassipoureaceaeCassythacea

e Castaneaceae Castelaceae Casuarinaceae Catesbaeaceae Cathedraceae Cecropiace

ae CedraceaeCedrelaceae Celastraceae Celosiaceae Celtidaceae Centaureaceae Cen

trolepidaceae Centroplacaceae CepaceaeCephalanthaceae Cephalotaceae Cephalota

xaceae Cerastiaceae Ceratoniaceae Ceratophyllaceae CerberaceaeCercidiphyllaceae

Cercocarpaceae Cercodiaceae Cereaceae Cerinthaceae Ceroxylaceae Cestraceae Cev

alliaceaeChailletiaceae Chamaedoraceae Chamaemoraceae Chamelauciaceae Chaun

ochitonaceae Chelidoniaceae ChelonaceaeChenopodiaceae Chimonanthaceae Chingi

thamnaceae Chionographidaceae Chiranthodendraceae ChironiaceaeChlaenaceae Ch

loanthaceae Chloranthaceae Chloridaceae Chlorogalaceae Chrysobalanaceae Cicerac

eae CichoriaceaeCimicifugaceae Cinchoniaceae Circaeaceae Circaeasteraceae Cissac

eae Cistaceae Citraceae Clematidaceae CleomaceaeClethraceae Clusiaceae Cneorac

eae Cnestidaceae Cnicaceae Cobaeaceae Cochlospermaceae Cocosaceae Coffeaceae

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Colchicaceae Coleogynaceae Columelliaceae Combretaceae Commelinaceae Comocl

adiaceae Compositae CompsoaceaeConantheraceae Connaraceae Conostylidaceae C

onvallariaceae Convolvulaceae Cordiaceae CoreopsidaceaeCoriandraceae Coriariacea

e Coridaceae Corispermaceae Cornaceae Corokiaceae Coronillaceae Correaceae Cor

rigiolaceaeCorsiaceae Corydalaceae Corylaceae Corynocarpaceae Coryphaceae Cost

aceae Cotyledonaceae CoulaceaeCoutariaceae Coutoubiaceae Crassulaceae Crescen

tiaceae Cressaceae Crinaceae Crocaceae CroomiaceaeCrossosomataceae Crotonacea

e Cruciferae Cryptaceae Crypteroniaceae Cryptocorynaceae CryptomeriaceaeCtenol

ophonaceae Cucurbitaceae Cunninghamiaceae Cunoniaceae Cupressaceae Curcumac

eae Curtisiaceae CuscutaceaeCyananthaceae Cyanastraceae Cynanchaceae Cyanella

ceae Cycadaceae Cyclanthaceae CyclantheraceaeCyclocheilaceae Cydoniaceae Cymo

doceaceae Cynaraceae Cynocrambaceae Cynomoriaceae Cyperaceae CyphiaceaeCyp

hocarpaceae Cypripediaceae Cyrillaceae Cyrtandraceae Cyrtanthaceae Cytinaceae

Dacrycarpaceae Dactylanthaceae Damasoniaceae Daphnaceae Daphniphyllaceae Das

ypogonaceae DatiscaceaeDaturaceae Daucaceae Davidiaceae Davidsoniaceae Decais

neaceae Deeringiaceae Degeneriaceae DelphiniaceaeDendrophthoaceae Columelliac

eae Detariaceae Dialypetalanthaceae Dianellaceae Dianthaceae DiapensiaceaeDicha

petalaceae Dichondraceae Diclidantheraceae Dicrastylidiaceae Dictamnaceae Didiere

aceae DidymelaceaeDidymocarpaceae Diegodendraceae Diervillaceae Digitalidaceae

e Dilatridaceae Dilleniaceae Dioaceae DionaeaceaeDioncophyllaceae Dioscoreaceae

Diosmaceae Diospyraceae Dipentodontaceae Diphylleiaceae Diplolaenaceae Dipsaca

ceaeDipterocarpaceae Dirachmaceae Disanthaceae Diselmaceae Dodonaeaceae Do

mbeyaceae Donatiaceae DorsteniaceaeDortmannaceae Doryanthaceae Drabaceae D

racaenaceae Dracontiaceae Drimyidaceae Droseraceae DrosophyllaceaeDryadaceae

Duabangaceae Duckeodendraceae Dulongiaceae Durantaceae Durionaceae Dysphani

aceae

Ebenaceae Eccremidaceae Ecdeiocoleaceae Echiaceae Echinopaceae Ehretiaceae Ela

eagnaceae ElaeocarpaceaeElegiaceae Elatinaceae Ellisiaceae Ellisiophyllaceae Elodea

ceae Elytranthaceae Embeliaceae Emblingiaceae EmmotaceaeEmpetraceae Encepha

lartaceae Engelhardtiaceae Enhalaceae Epacridaceae Ephedraceae Ephemeranthacea

eEpilobiaceae Epimediaceae Eragrostidaceae Eremolepidaceae Eremosynaceae Erica

ceae Erinaceae EriocaulaceaeEriogonaceae Eriospermaceae Erodiaceae Erycibaceae

Eryngiaceae Erysimaceae Erythroniaceae ErythropalaceaeErythrospermaceae Erythro

xylaceae Escalloniaceae Eschscholziaceae Eucomidaceae Eucommiaceae Eucryphiace

aeEuonymaceae Eupatoriaceae Euphorbiaceae Euphrasiaceae Euphroniaceae Eupo

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matiaceae Eupteleaceae EuryalaceaeEustrephaceae Euthemidaceae Eutocaceae Exb

ucklandiaceae Exocarpaceae

Fabaceae Fagaceae Falcatifoliaceae Ferulaceae Festucaceae Ficaceae Ficoidaceae Fit

zroyaceae FlacourtiaceaeFlagellariaceae Flindersiaceae Foetidiaceae Forestieraceae

Fothergillaceae Fouquieriaceae Fragariaceae FrancoaceaeFrangulaceae Frankeniacea

e Fraxinaceae Fraxinellaceae Freycinetiaceae Fritillariaceae Fuchsiaceae Fumariacea

eFunkiaceae

Gaiadendraceae Galacaceae Galanthaceae Galaxiaceae Galedupaceae Galeniaceae G

aliaceae Garciniaceae GardeniaceaeGardneriaceae Garryaceae Geissolomataceae Ge

itonoplesiaceae Gelsemiaceae Geniostomaceae GentianaceaeGeonomataceae Geosir

idaceae Geraniaceae Gerrardinaceae Gesneriaceae Gethyllidaceae Gilliesiaceae Gina

lloaceaeGinkgoaceae Gisekiaceae Gladiolaceae Glaucidiaceae Glechomaceae Glinace

ae Globulariaceae Gnaphaliaceae GnetaceaeGoetzeaceae Gomortegaceae Gomphiac

eae Gomphrenaceae Gonystylaceae Goodeniaceae Gordoniaceae GouaniaceaeGoup

iaceae Gramineae Gratiolaceae Grewiaceae Greyiaceae Grielaceae Griseliniaceae Gr

onoviaceae GrossulariaceaeGrubbiaceae Guaiacanaceae Guamatelaceae Guettardac

eae Gunneraceae Gustaviaceae Guttiferae GyrocarpaceaeGyrostemonaceae

Hachetteaceae Haemanthaceae Haemodoraceae Halesiaceae Hallieraceae Halocarpa

ceae Halophilaceae HalophytaceaeHaloragaceae Hamamelidaceae Hanguanaceae H

aptanthaceae Harmandaceae Hebenstretiaceae HectorellaceaeHederaceae Hedyosm

aceae Hedyotidaceae Hedysaraceae Heisteriaceae Heleniaceae Heliamphoraceae H

elianthaceaeHelianthemaceae Helichrysaceae Heliconiaceae Helicteraceae Heliotropi

aceae Helleboraceae Heloniadaceae HelosidaceaeHelwingiaceae Hemerocallidaceae

Hemimeridaceae Hemiphylacaceae Henriqueziaceae Henslowiaceae Hermanniaceae

Hernandiaceae Herniariaceae Herreriaceae Hesperocallidaceae Heterantheraceae He

teropyxidaceae HeterostylaceaeHibbertiaceae Hibiscaceae Hilleriaceae Himantandra

ceae Hippocastanaceae Hippocrateaceae HippomanaceaeHippophaeaceae Hippurida

ceae Hirtellaceae Holacanthaceae Homaliaceae Hopkinsiaceae Hoplestigmataceae H

ordeaceaeHornschuchiaceae Hortensiaceae Hortoniaceae Hostaceae Houstoniaceae

Huaceae Huerteaceae HugoniaceaeHumbertiaceae Humiriaceae Hyacinthaceae Hyd

atellaceae Hydnoraceae Hydrangeaceae Hydrastidaceae HydrillaceaeHydrocharitacea

e Hydrocotylaceae Hydrogetonaceae Hydroleaceae Hydropeltidaceae Hydrophylacac

eae HydrophyllaceaeHydrostachyaceae Hymenocardiaceae Hypecoaceae Hyperanth

eraceae Hypericaceae Hypopityaceae HypoxidaceaeHypseocharitaceae

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Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta

ceae ImperatoriaceaeInocarpaceae Inulaceae Ionidiaceae Iriarteaceae Iridaceae Irvi

ngiaceae Isnardiaceae Isophysidaceae IteaceaeStrasburgeriaceae Ixiaceae Ixioliriace

ae Ixonanthaceae

Jalapaceae Jamboliferaceae Japonoliriaceae Jasionaceae Jasminiaceae Johnsoniaceae

Joinvilleaceae JuglandaceaeJulianiaceae Juncaceae Juncaginaceae Juniperaceae Jus

siaeaceae Justiciaceae

Kadsuraceae Kaliphoraceae Kalmiaceae Kaniaceae Kiggelariaceae Kingdoniaceae Kin

giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae

Krameriaceae

Labiatae Lacandoniaceae Lachenaliaceae Lacistemataceae Lactoridaceae Lactucaceae

Lagerstroemiaceaee LamiaceaeLampsanaceae Lanariaceae Langsdorffiaceae Lantan

aceae Lapageriaceae Lardizabalaceae Lasiopetalaceae LauraceaeLawsoniaceae Laxm

anniaceae Lecythidaceae Ledaceae Ledocarpaceae Leeaceae Leguminosae Leitneriac

eae LemnaceaeLennoaceae Lentibulariaceae Lentiscaceae Leoniaceae Leonticaceae

Lepidobotryaceae LepidocarpaceaeLepidocaryaceae Lepidocerataceae Leptaulaceae

Leptospermaceae Lepturaceae Lepuropetalaceae LeuctenbergiaceaeLibocedraceae

Licaniaceae Ligustraceae Lilaeaceae Liliaceae Limeaceae Limnanthaceae Limnocharit

aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace

ae Linderniaceae Lindleyaceae Linnaeaceae LippayaceaeLiriaceae Liriodendraceae Li

ssocarpaceae Littorellaceae Loasaceae Lobeliaceaee Loganiaceae LomandraceaeLoni

ceraceae Lopeziaceae Lophiocarpaceae Lophiolaceae Lophiraceae Lophophytaceae L

ophopyxidaceae LoranthaceaeLotaceae Lowiaceae Lupulaceaee Luxemburgiaceae Lu

zuriagaceae Lychnidaceae Lyciaceae LyginiaceaeLygodisodeaceae Lysimachiaceae Ly

thraceae

Macarisiaceae Mackinlayaceae Madiaceae Maesaceae Magnoliaceae Malaceae Male

sherbiaceae MalortieaceaeMalpighiaceae Malvaceae Mangiaceae Manicariaceae Ma

paniaceae Marantaceae Marathraceae MarcgraviaceaeMartyniaceae Mastixiaceae

Matricariaceae Maundiaceae Mayacaceae Medeolaceae Medusagynaceae Peridiscac

eaeMelaleucaceae Melampyraceae Melanophyllaceae Melanthiaceae Melastomatac

eae Meliaceae Melianthaceae MelicaceaeMeliosmaceae Melittidaceae Melochiacea

e Memecylaceae Mendonciaceae Menispermaceae Menthaceae MenyanthaceaeMe

nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

47

Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

Nypaceae Nyssaceae

Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

ceae Papilionaceae Pappophoraceae PapyraceaeParacryphiaceae Parasitaxaceae Par

ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

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hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

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cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

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ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 5: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

5

somehow reflects the degree of morphological differences between taxa or that taxa at

the same rank be based on similar characters or show a similar amount of distinctness

This might be possible using phenetic methods of analysis but is very difficult if ones

classification is phylogeny-based as here it could promote instability if used in taxa

where such a criterion had not previously been used and it might also inadvertently

suggest that taxa might be equivalent (for an example see Fritsch et al 2008) There

have also been proposals that rank could reflect the age of the clade (eg Hennig 1966)

with clades that have diverged by a particular time all being given the same rank Apart

from the fact that aging times of divergence of clades is still a difficult enterprise huge

disruptions to our nomenclature would result Recent suggestions which invoke the use

of age in classifications focus on providing a standardized timeclip ie a set of letters

referring to a particular geological period that could simply be added to a conventional

taxon name (Avise amp Mitchell 2007) However even such timeclips are unlikely to come

into general use soon

It would be impossible even to think about a higher-level classification such as this

without the advances in our understanding of relationships made by the phylogenetic

analyses of molecular data carried out over the last twenty years One can then

integrate the data to be found in both classical and recent morphological studies with

these phylogenies For the dramatic changes in this area see for instance the

pessimistic attitude towards orders in Davis and Heywood (1963 107-108) The most

unsatisfactory taxon in Angiosperm classification they were indefinable their

circumscription was not fixed etc Families they thought were likely to be the largest

natural unit within the mono- or dicotyledons Along the same lines almost three

quarters of the orders (4459 monofamilial orders ignored) recognised by Cronquist

(1981) are not monophyletic ie they do not contain all and only the descendents of a

common ancestor Most of those orders that are monophyletic are very small

(Zingiberales with eight families are the largest) for families on the other hand about

two thirds (189273) are monophyletic

Turning now to phylogenetic classifications and the particular classification used

here Backlund and Bremer (1998) provide a useful discussion on the principles of

phylogenetic classification that is applicable at all levels apart from species (see also

Stevens 1998 also Albach et al 2004 Entwisle amp Weston 2005 Pfeil amp Crisp 2005 etc

for examples) Backlund and Bremers main principle is that taxa that are recognised

formally should be monophyletic However this does not indicate which particular

clades we might wish to name as families genera etc and talk about in general

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

6

conversation If a well-supported hypothesis of monophyly is a necessary prerequisite

for a group to be named it is not a sufficient prerequisite (but cf the PhyloCode -

Cantino amp de Queiroz 2006 Cantino et al 2007) Not all clades need be named indeed

it would barely be practicable (or practical) to do this To decide which clades should be

named additional criteria can be invoked Other things being equal it is helpful if 1 taxa

formally recognised are easily recognizable 2 groups that are well-established in the

literature are preserved 3 the size of groups is taken into account and 4

nomenclatural changes are minimized (Backlund amp Bremer 1998) Thus numerous small

groups have little to recommend them since individually they summarize little

information and tend to clog the memory while groups that are too big may be

amorphous Somewhat similarly Godfray and Knapp (2004 p 562) note that users

want stable informative and accessible classifications that enable easy identification

(see also Simon 2008) - although invoking users without specifying those who make up

this group is not very helpful This classification is for all interested in comparative

biology hence the emphasis on monophyly Although the clades named are sometimes

difficult to characterise there are as we shall see many ways of making such a

classification accessible to all

Problems with this emphasis on monophyly may be caused by reticulation events

such as hybridization endosymbiosis and lateral gene transfer but they are unlikely to

be common confusing factors here Genera can often be pegged to above the level at

which hybridization is at all common However in Poaceae-Pooideae-Triticeae there are

some intractable problems where extremely well established common usage and the

principle of monophyly are likely to remain at odds Many genera are certainly not

monophyletic here being allopolyploids and the genera are ultimately based on the

different genomes they contain (Dewey 1984 Loumlve 1984 Barkworth 2000 for a history

of Triticeae classification Petersen et al 2006) There is also extensive reticulation

reported within Danthonioideae (Pirie et al 2009) Evidence also increases of old

hybridization events elsewhere in flowering plants that at the very least cause

discordance between relationships suggested by different genomic compartments as in

Smedmark and Anderberg (2007 Sapotaceae) and Fehrer et al (2007) Morgan et al

(2009) and Pelser et al (2008 all Asteraceae - hybridization is likely to be quite a

problem at the generic level here) and genera like Medicagoare turning out to have

highly reticulating relationships at the species level (Maureira-Butler et al 2008) Of

course there are many problems trying to apply the concept of monophyly to species

and for many - but not all - biologists strict monophyly is a less important criterion at

this level (Funk amp Omland 2003 see much of the discussion in Houmlrandl 2006)

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

7

The major endosymbiotic events that characterize the clade of which flowering

plants are a part (and gave rise to chloroplasts and mitochondria) are very ancient and

cause no problems for the student of multicellular organisms However lateral gene

transfer has been detected in a number of situations between quite unrelated

organisms (eg Bergthorsson et al 2003 - Amborella and liverworts see also below but

cf Goremykin et al 2009) and it may be particularly common in mitochondria (Sanchez-

Puerta et al 2008) Here too there are no major problems providing one is careful

such transfers do however raise all sorts of interesting biological questions (see

Richardson amp Palmer 2007 for a summary) Although there is increasing evidence for the

importance of genome duplications - hybridization is one cause of this - at various times

during the evolution of seed plants and of palaeopolyploidy events within eg the

Lauraceae and Magnoliaceae clades (Soltis et al 2009 for a summary see eg

the Characters page for further discussion) these too do not currently seem to pose

problems for the adoption of monophyly as the sine qua non of groups to be recognised

formally in this phylogenetic classification but it can make detecting orthologous genes

difficult

The accessory principles of Backlund and Bremer (1998) should be used in

combination Thus keeping the monogenericPlatanaceae separate from its sister

taxon Proteaceae is justifiable Both are much-used names that signal well supported

well defined and easily recognisable groups that have long been recognised as distinct

have several synapomorphies and do indeed look very unlike each other Combining

the two would yield a clade with few obvious apomorphies not to mention the fact

that Nelumbonaceae should by the same logic (it is also monogeneric) also be included

in the expanded family On the other hand it is difficult to justify the continued

recognition of Callitrichaceae or Hippuridaceae monophyletic and distinctive although

they may be If they were recognised several poorly characterised clades would also

have to be carved out ofPlantaginaceae in any classification that aimed to convey a

comprehensive view of the worlds flora The continued recognition

of Valerianaceae and Dipsacaceae also tends to run into this problem (see also Pfeil amp

Crisp 2005 Orthia et al 2005 Albach 2008 etc for useful practical discussions of such

matters) But there are no absolute guidelines If Podostemaceae turn out to be sister

to Hypericaceae (for references here and elsewhere in the Introduction see the

individual families) the subsequent moderate dismemberment of Clusiaceae sl is not

be too high a price to pay for the continued recognition of Podostemaceae Hence the

somewhat provisional recognition of Hypericaceae and Calophyllaceae as well as

Clusiaceae below the families can all be recognized and the name Podostemaceae in

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

8

particular is very well established In a somewhat similar situation if Lemna and its

relatives are a clade sister to most other Araceae should they be recognised as a

separate family Gymnostachys a phenetically fairly distinctive taxon as well as the less

phenetically distinct Orontioideae would have to be recognised as a separate families

(or combined as a single family) too but Araceae in a somewhat restricted sense would

be somewhat more morphologically coherent although not greatly so and not notably

distinct However it is in the very nature of such decisions to be somewhat arbitrary

and unsatisfactory hence the emphasis on consensus classifications here and on

classifications as simply being a means to an end

A useful distinction can be drawn between crown groups and stem groups The

former are monophyletic and include the extant members of a clade and their

immediate common ancestor (see the figure below) The groups characterized in this

site are such groups Thus Proteaceae here are crown group Proteaceae apomorphies

like the single carpel four-merous perianth etc being found in this common ancestor

Stem groups on the other hand include all the members of a lineage immediately after

its split from its sister group and all branches of this lineage In the case of Proteaceae

the stem group would include everything after its split from its sister group

Platanaceae Obviously most of the organisms in the lineage that terminates in crown-

group Proteaceae are unknown only a few being known as fossils and it is not known

exactly where particular apomorphies of crown group Proteaceae evolved along this

lineage

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

9

Indeed there is no reason other than convention or convenience why any group

should not be segregated into several smaller monophyletic groups or merged to

produce a larger unit we can talk about one large thing or about several smaller things

Thinking about aspects of size findings in ethnobiology and cognitive psychology can be

used to suggest that a moderate number - probably fewer than 500 - of families is a

reasonable goal at which to aim and that groupings of taxa throughout any system

should be rather small in size (eg Berlin 1992 Stevens 1994 1997) Major systems such

as those of Linnaeus and Bentham and Hooker were constructed explicitly so as to ease

the burden on the memory (Stevens 1997 2002 see also Scharf 2007) the latter in

particular ensuring that all groups in their classification were relatively small often

containing three to eight immediately subordinate taxa - but by no means all their

groups were formally named Along the same lines Burtt (1977b) suggested that the

number of names at any rank should be at most one third those at the immediately

lower rank - and monotypic taxa might not need a formal name Consistent with such

ideas a fairly broad view of families and orders is taken here whenever the constraints

of monophyly and other criteria used when constructing classifications (see above)

permit The APG system is thus a convention consistent with ideas such as those

expressed by Backlund and Bremer (1998) for similar conventions see eg the Grass

Phylogeny Working Group (2001) and for fungi Hibbett et al (2007)

There are other views Thus Takhtajan (1997) has suggested that smaller families

are more natural This is incorrect Monophyletic groups that include fewer taxa -

Takhtajans smaller families - do not necessarily have more apomorphies than larger

groups even if members of smaller groups will always have more features in general in

common than the larger group that includes them That is they will have their

apomorphies their unique features as well as progressively more plesiomorphies

features found both in the small groups and in the larger clades of which they are a part

having more features in common is indeed one common meaning of more natural

(Note that the implication of the word natural has long been a group of the kind

[usually unspecified] that I think should be recognised and if other botanists had

different ideas too bad their groups were necessarily not natural [see Bather 1927] It

is thus rarely a helpful word and is not used here) Furthermore if this approach is

adopted we will find a slippery slope ahead By this kind of argument all families should

be very small since their members will have a great deal in common and so will be

most natural However as families (for example) are split the relationships that are

evident between the segregates and that were responsible for their being placed in a

single family in the first place will seem to necessitate the recognition of a new order

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

10

etc as is evident in Takhtajans own work - general taxonomic inflation is the result (see

also comparable suggestions in a cladistic context for Brassicales in particular - Ronse de

Craene amp Haston 2006) Such splitting is also questionable when teaching and learning

families since the student needs to understand the system as a whole However for

some genera removed from the families that until now have included them the

phenetic-classificatory-phylogenetic structure in their new home may mandate the

recognition of small families On the other hand Takhtajans suggestion that narrowly

defined families are more useful for phylogenetic studies may be true Indeed I have

more than once regretted prematurely combining groups whether species (in the

context of monographic work) or families (in the course of preparing these notes) but

this is largely a separate issue

Van Steenis (1978) Philipson (1987b) and others have rightly questioned the value

of splitting a group when ideas of the relationships of its constituent members have not

changed - that is very good reasons have to be provided for splitting a family if the

genera within it remain part of the same clade rather than belonging to another clade

Thus APG (2003) broadened the circumscription of Malvaceae because of the

parapolyphyly of some of the families that had historically been associated with it (Judd

amp Manchester 1997 Alverson et al 1999 Bayer et al 1999) These families particularly

Tiliaceae and Sterculiaceae were not at all easy to distinguish their close relationship

(see eg Brown 1814) and overall similarity had long been conceded and to some

workers at least their combination has come as something of a relief Although most of

the larger clades within Malvaceae sl remain difficult to distinguish even with flowers

Cheek (2007) opts for a wholesale and novel dismemberment into ten families

however the very good reasons for doing this are wanting

The same principles are of course applicable when it comes to dividing genera little

other than a headache is gained by splitting genera such

as Drosera and Gnetum (Doweld 2000) as has recently been proposed Thus if an

established genus divides into two (or more) clades this is not a signal for recognising

two groups at the same level - so here it could be argued that the dismemberment

of Pterostylis (Jones amp Clements 2002b) was somewhat unfortunate Along the same

lines if a newly-discovered taxon is sister to an existing named taxon - say a genus - this

does not necessitate the description of a separate genus for the newly described species

(cf Davis 2002) (Of course some [eg Thorne 1976] have suggested that the sizes of

gaps between groups at the same rank should be similar but any principle like this is

inherently flawed since morphological gaps are more unstable than phylogenetic

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

11

relationships and applying it across all flowering plants would both be difficult and cause

substantial changes - see also below) Humphreys and Linder (2009) provide a well-

documented survey of generic concepts in plants which the reader should consult they

note that generic limits (broad versus narrow) have oscillated historically and that

currently larger genera tend to be recognised because studies tend to be on a broader

scale than in the past

Note that invoking similarity or difference - whether qualified (considerable

similarities substantial differences) or not - in a cladistic context as justification for

combining or splitting taxa is not a particularly strong argument (see eg

CardiopteridaceaeStemonuraceae - Karingrehed 2002c) Similarity and difference can

neither be defined precisely since what may seem to be substantial similarities to me

may not to the next person nor are they likely to be stable in the face of our changing

knowledge of morphology and what might be synapomorphies

I might have prefered to merge some families recognised here or split others but

by and large I do not think my own preferences matter very much - and I take the same

position with regards to comparable preferences expressed by others Indeed the

bottom line is that in flagged hierarchies of the kind used here the limits of any

monophyletic unit generally taught and discussed particularly other than species can

be established only by convention and consensus (eg Stevens 2002 2006a Entwisle amp

Weston 2005) This is rather different from the reasonable observation that the

phylogeny itself cannot be achieved by consensus (Thorne 1976) for in most cases there

is a fact of the matter when it comes to relationships Given the increasing support for

the outlines of angiosperm phylogeny a stable consensus classification based on this

phylogeny seems attainable Indeed in addition to providing current ideas of

relationships of seed plants in a synthesised form this site is part of an attempt to build

such a consensus about the circumscription of taxa (see APG 1999 2003 2009 Grass

Phylogeny Working Group 2001 Mabberley 2008) Reaching such a consensus is vital

since what we know of angiosperm phylogeny allows a very large number of

classifications to be based on it and as we find out more the number of possible

classifications increases greatly Unfortunately however nature does not dictate what

the classification should be All classifications are constructed by humans to

communicate particular aspects of groups and relationships Our goals as systematists

are surely to produce robust hypotheses of relationships to understand the evolution of

morphology and the like - but not to argue ad nauseam whether something should be

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

12

a family or a subfamily That way surely lies madness and worse the discredit of our

discipline

There are similar issues whatever naming system is used Thus in phylogenetic

naming (Baum et al 1998 for an example but cf Baum et al 2004 for the PhyloCode

see Cantino amp De Queiroz 2006) an unflagged hierarchy is used in which any

terminations of names used are uninformative about the relative position of taxa If one

adopts the principle of phylogenetic naming one indeed does not have to worry about

the nomenclatural consequences caused by lumping or splitting any well-supported

clade can be named without affecting the name of more or less inclusive clades

Unfortunately unflagged hierarchies have very serious deficiences as communication

devices because they lack one aspect essential in language biological or otherwise -

they contain no intrinsic information about the relationships of the group in question to

others (eg Pfeil amp Crisp 2005 Stevens 2006a) Recent suggestions for using prefixes like

Apo- and Pan- to PhyloCode names will however allow limited information of this

kind to be conveyed but only as it pertains to individual branches and current

proposals do not even mandate that the prefixes be employed consistently In any

event such proposals simply prevent the potential tripling of the number of quite

different names used to describe different aspects of a phylogenetic tree over those

used to name monophyletic groups pure and simple In general where n is the number

of extant species in a group the number of clades in such a group = n-1 (Species will

also need names too for their names see Dayrat et al 2008) Importantly here too

consensus over the clade names commonly learned by students and used in herbaria is

needed otherwise communication will be impeded the names themselves will provide

no guidelines as to which should be chosen The situation is of course more complicated

than this Terminations that convey ideas of rank in a phylogenetic classification can also

be used in phylocode names - however there they will carry no implications of rank

How they will be used is another matter of course

Of course there are other philosophies of classification and some still prefer

evolutionary classifications There classificatory principles differ substantially from those

followed here eg the recognition of paraphyletic taxa may be permitted however

detailed reasons for prefering the taxa that are recognised are rarely given although

nature and natural groups are often mentioned (cf Stuessy amp Koumlnig 2008) For

summaries of commonly used systems see Brummitt (1992) and Mabberley (2008)

new evolutionary systems appear every year or so However even those who allow or

promote the recognition of paraphyletic groups (eg Grant 2003 Thorne 2007

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

13

Heywood et al 2007) may find it of interest to examine a system recognizing only

monophyletic groups it provides a rather different understanding of evolution

Evolutionary classifications in general try and combine phylogeny and

morphological gaps although that is no easy thing to do - it is akin to combining chalk

and cheese (for an attempt to make this impossible task seem to be more objective see

Stuessy amp Koumlnig 2008) However there is a principle from evolutionary classification

that is relevant and I think quite useful here The size of the gap between two groups

tends to be inversely proportional to the sizes of the groups involved (Davis amp Heywood

1963) One can imagine a situation in which a large group is formally divided even

although the distinguishing characters of the two are weak whereas a smaller group

similarly divisable will be left intact

To summarize If hypotheses of phylogeny remain stable we should be able to base

a stable classification on that phylogeny and then get on with our work that is testing

the phylogenies we have elucidating phylogenies in areas where relationships are

unclear studying the evolution of morphology describing species etc In this context

the spread of the Angiosperm Phylogeny Group system (see below) and its widespread

utilisation in technical literature also floras (eg van der Meijden 2005) textbooks (eg

Simpson 2006 Judd et al 2007 [third edition]) dictionaries (Mabberley 2008) more

popular literature (eg Souza amp Lorenzi 2005 Spears 2006) and as an outline for a new

herbarium sequence (Haston et al 2007) is gratifying The posibility that one might be

able to develop a stable phylogeny-based classification of families and in particular

orders represents a dramatic turn-around from the pessimistic attitude about such

higher-level groupings expressed by Davis and Heywood (1963) and Thorne (1976) the

latter even suggesting that we should bury forever the metaphor of the phylogenetic

tree as highly unrealistic (ibid p 56) Returning to Godfray and Knapps (2004) users

of classifications who want a stable informative and accessible classification that

enables easy identification - unfortunately they want cake with everything and cannot

get it - these pages attempt to satisfy as many of their needs as possible but phylogeny

and monophyly are the primary shapers of the APG classification

On this classification in particular

Here I very largely follow the Angiosperm Phylogeny Group classification (APG

2003) Any differences are not to be interpreted as differences in principle simply that

new phylogenies continue to be published and here I attempt to provide an overview

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

14

of current ideas of higher-level relationships of seed plants The Angiosperm Phylogeny

Group classification is based on well-supported relationships evident in the numerous

molecular studies that began to appear in the late 1980s (see APG 1999 for the

principles underlying the classification) but recent phylogenetic work does not

contradict the major outlines of the trees used by APG II (2003) or even those in APG

I (APG 1999) The APG classification has for the most part been conservative and I

am similarly conservative here For the names and authorities I follow APG III (2009)

although the names of the authors may not always be correct these being in something

of a state of flux for the superordinal names used see Chase and Reveal (2009)

although classification at this level here is not exhaustive - only clades with several

orders have superordinal names

Some changes in our ideas of relationships and hence in the clades we talk about

are particularly likely in parts of Lamiales Caryophyllales and Malpighiales for example

although this will not affect the orders themselves Thus some changes to clade

circumscriptions are to be expected although I do not expect them to be substantial

but changes are neither a defect of cladistics nor a necessary consequence of the use of

molecular data Clades are hypotheses of relationships and as hypotheses they may be

overturned However work since APG I has not suggested other than minor changes

in the compositions of the orders even if the odd genus or even family is turning out to

be seriously misplaced - recent examples are Hydatellaceae (from monocots-Poales to

Nymphaeales Saarela et al 2007) Guamatelaceae (from Rosales-Rosaceae to

Crossosomatales Oh amp Potter 2006) and Perrottetia and Bhesa (from Celastrales-

Celastraceae to Huerteales and Malpighiales respectively Zhang amp Simmons 2006) The

main changes have been clarification of the relationhips of individual families or groups

of families that were of uncertain position eg of Chloranthaceae (Moore et al 2007)

Ceratophyllaceae (Jansen et al 2007) and unplaced asterid II families (Winkworth et al

2008a) As mentioned above phylogenies in the area of main interest in these pages are

overwhelmingly tree-like Although there is evidence for acquisition of host plant genes

by parasites (see below) even wider but inexplicable transfer of mitochochondrial

genes in plants like Amborella (Bergthorsson et al 2004 cf Goremykin et al 2009a)

and even transfers of nuclear genes (Vallenback et al 2008) these are the exception

rather than the rule

In cases where the Angiosperm Phylogeny Group II allowed alternatives as to the

limits of families - Papaveraceae in the broad sense or Papaveraceae plus

Pteridophyllaceae plus Fumariaceae Proteaceae in the broad sense or Proteaceae plus

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

15

Platanaceae - the choices made here follow common usage eg as in textbooks like

Judd et al (2007) and Simpson (2006) and particularly in the new edition of

Mabberleys The Plant Book (Mabberley 2008) This latter is itself an attempt to reflect a

consensus the result of taking the opinions of botanists at several meetings A largely

similar consensus is reflected in the APG III classification (APG 2009) where you will

see that alternative classifications have been dispensed with For many people the

existence of such alternative classifications will simply confuse so agreement over

which groupings to use when alternatives are permitted and then dispensing with the

alternatives that are not commonly used seems reasonable I am making the few

changes that are needed to bring the classification here in line with these others but

only gradually since there is not enough time in each day

As already mentioned higher-level relationships in general and the composition of

orders in particular have always presented something of a challenge to systematists

(eg Davis amp Heywood 1963) That being said the composition of some clades like

Apiales Crossosomatales and Pandanales is decidedly unexpected However it is

interesting to see that these higher level clades are generally accepted even in works

with different classificatory philosophies (eg Heywood et al 2007) For clades like

Malpighiales many of the family groupings within Asparagales etc attempts to find

distinctive characters have largely failed (but see Endress amp Matthews 2006a also

the Apomorphies page here) Interestingly as with families some groupings suggested

by molecular studies are supported by morphological andor chemical characters that

have long been known sometimes for over a hundred years the relationship between

Pittosporaceae and ApiaceaeAraliaceae are a case in point (Hegnauer 1969b and

references) As our knowledge of morphology and chemistry improves we can hope for

improvements in the characterisations of clades at all levels

In many cases the new family limits of the Angiosperm Phylogeny Group (see

APG 1999 2003) are not really controversial although changes from the limits

commonly accepted only a decade ago are sometimes quite dramatic (eg Wagenitz

1997) Thus the split of the old Saxifragaceae sl is necessitated by its extreme

polyphyly as also with Icacinaceaesl and Cornaceae sl However the limits and

relationships of such groups had long been considered to be unsatisfactory but until

recently there had been no real reason to prefer one proposed arrangement over

another It is generally accepted that the limits of Lamiaceae and Verbenaceae have to

be redrawn although there is as yet no compelling evidence that the redrawn taxa are

not sister taxa (there is no evidence that they are) But whatever their relationships the

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

16

content of the clades has changed considerably and incidentally they are now easier to

identify than before the decision to recognise two families is not difficult The same is

true for Salicaceae and Achariaceae (Malpighiales) two previously small families that

have received the bulk of the old Flacourtiaceae Clade and hence taxon limits remain

difficult around Theaceae (Ericalessee alsoSladeniaceae Pentaphylacaceae)

and Euphorbiaceae (Malpighiales see

also Phyllanthaceae Picrodendraceae Putranjivaceae etc) Here current groupings

may not represent quite such dramatic changes in our understanding of relationships

Indeed parts of the old Euphorbiaceae that were separated may yet go back together

particularly Phyllanthaceae and Picrodendraceae (see Wurdack et al 2004 Davis et al

2005) even if Theaceae should indeed be dismembered (Geuten et al 2004) However

given that there is no molecular evidence that currently warrants combining all the

segregates of Euphorbiaceae and even if some do come together the clades the

families represent suggest novel groupings not recognised in current classifications

Furthermore Rafflesiaceae appear to be embedded within Euphorbiaceae s str (Davis

et al 2007) so the family is divided Of course maintaining even a moderately narrowly-

circumscribed Euphorbiaceae would entail reducing the iconic Rafflesiaceae to

synonymy Relationships in core Caryophyllales especially

around Phytolaccaceae andNyctaginaceae and also Portulacaceae are incompletely

understood and refashioning of taxon limits will doubtless be needed as cladistic

relationships become apparent (eg see Nyffeler 2007 Ogburn 2007a b) Some

groupings in the old Icacinaceaeand Olacaceae also remain unclear

The discovery of the relationships of parasitic and aquatic groups have presented a

particular challenge to systematists Morphologically some of these plants are so highly

modified that interpretation of the plant body in conventional terms is difficult or even

impossible Thus parasitic groups (broad-leaved angiosperms only) such as Rafflesiaceae

are hard to place since both the vegetative body and the flowers are changed almost

beyond recognition (flowers of rafflesiaceae are a good example of gigantism - Davis et

al 2007) Furthermore plastid gene sequences may be difficult or impossible to obtain

the chloroplast DNA in particular being highly degraded and the rate of molecular

change in general may be high so the problem of long-branch attraction is serious (eg

Duff amp Nickrent 1997 Nickrent et al 1998 Caddick et al 2002a G Petersen et al

2006b) Echlorophyllous sapromycoheterotrophic taxa mostly known from monocots

present similar problems However progress is being made placements for

Apodanthaceae Rafflesiaceae Mitrastemonaceae Cytinaceae and Cynomoriaceae have

recently been suggested (Barkman et al 2004 2007 Davis amp Wurdack 2004 Nickrent et

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

17

al 2004 Davis et al 2007) it seems likely that Burmanniaceae sl are polyphyletic but

both parts are in Dioscoreales (Merckx et al 2006) and relationships within the largely

hemiparasitic Santalales are also gradually being clarified (Maleacutecot 2002) The inclusion

of parasitic taxa in general molecular analyses can cause conniptions (eg Nickrent et al

2004 Davis et al 2004 Chase et al 2006 G Petersen et al 2006b) and there can be

horizontal transmission of genes (eg Davis amp Wurdack 2005 Vitaceae to Rafflesiaceae

Barkman et al 2007 the mitochondrial atp1 gene commonly moves)

Similarly in water plants neither vessels in particular nor much xylem in general is

needed leaves are highly modified and water-mediated pollination if adopted may

well be associated with major changes in floral morphology Here too recent molecular

studies suggest that aquatic groups with hitherto problematic relationships may find

homes Thus Podostemaceae are close to Clusiaceae and Calophyllaceae and are sister

to Hypericaceae (Malpighiales Kita amp Kato 2001) Hydatellaceae which used to be in

Poales are part of Nymphaeales (Saarela et al 2007) and Hydrostachyaceae may be

close to Hydrangeaceae (Cornales Xiang et al 2002) or perhaps should be placed in

Lamiales note that in the first two cases in particular there are morphological and

chemical features that support such a move If Podostemaceae are indeed close to

Clusiaceae I look forward to seeing hypotheses to explain how the dramatic changes in

the vegetative body that have made Podostemaceae so problematic for generations of

systematists took place That conventional wisdom has trouble in understanding or

explaining how the morphologies of groups like Clusiaceae and Podostemaceae can be

related is largely a problem with conventional wisdom Ceratophyllaceae seem to be

finding a position as sister to eudicots (see Moore et al 2007) but their morphology is

so derived that there is no morphological evidence of which I am aware for this

relationship There are also similar although less extreme morphological problems with

many plants pollinated by wind

In such situations especially with some water plants mycoheterotrophs and

parasites a variety of characters may be affected and particularly under such

circumstances groups have tended to be formed using a few or even only a single

character that seems to provide evidence of relationships Add to this the tendency to

weight some characters particularly strongly on a priori grounds and the result was

either the recognition of conglomerate taxa such as Amentiferae which now appear to

be highly polyphyletic or the segregation of families like Plantaginaceae s str (now

much expanded) and Leitneriaceae (now in Simaroubaceae) - all of these are more or

less wind pollinated

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

18

Even although relationships of many parasitic and aquatic groups are being

clarified and at least some of the proposed phylogenetic positions seem to be well

supported it is indeed often still very difficult to understand how such groups have

evolved from their more morphologically conventional relatives However we have had

all sorts of preconceptions as to how characters should evolve Thus we used to

assume that features like highly scalariform vessel perforation plates or the complete

absence of vessels or a flower with an androecium that had many stamens a superior

ovary or separate petals were necessarily plesiomorphic or primitive and conversely

a simple perforation plate and the presence of vessels an androecium with few

stamens inferior ovary or petals that were connnate were almost necessarily

apomorphic or advanced (but cf eg Stebbins 1951) Such assumptions are incorrect

(eg Soltis et al 2005b) Carpels may become secondarily free carpels may fail to close

the seeds then developing outside the confines of the carpel as in some Aspagaraceae-

Nolinoideae Violaceae Berberidaceae Malvaceae-Sterculioideae etc) in Peliosanthes

teta perhaps the only species in Peliosanthes(Asparagaceae-Nolinoideae) the ovary

varies from superior to inferior (Jessop 1976 species limits here need close

investigation see also Kuzoff et al 2001 and Soltis amp Hufford 2002 Saxifragaceae

Apiales etc) many-seeded carpels can evolve from few-seeded carpels

(Razafimandimbison et al 2008) monoecy may be derived from dioecy (Schaefer amp

Renner 2010 and references) Classic studies such as those by Babcock (eg 1947)

on Crepis that assumed that evolution - in this case of the karyotype in particular - was

unidirectional have needed comprehensive re-evaluation (Enke amp Gemeinholzer 2008)

Most if not all characters have reversed andor evolved in parallel as is clear in the

discussion of ovary position here especially in Poales Saxifragales and Asparagales and

also in comments on such morphologically distinctive taxa (when compared with their

immediate relatives) such as Menyanthaceae and especially Pittosporaceae Parallel

evolution may occur even at the level of amino acid substitution as in the independent

acquisition of the phosphoenolpyruvate carboxylase (pepC) gene in C4 photosynthesis in

grasses (Christin et al 2007b see also Blaumlsing et al 2000)

Thinking of how such characters have been used in the past in classifications at the

generic level the results have been similar to those just discussed above for parasitic

plants and others Relying too much on animal pollination syndromes to mark generic

boundaries has all too often led to taxa that are highly unsatisfactory phylogenetically

(see eg Acanthaceae Bignoniaceae Campanulaceae Ericaceae Lamiaceae

Orchidaceae) and over-reliance on characters of fruit and seed (see eg Brassicaceae

and Apiaceae) has also led to unsatisfactory generic limits Again the more general

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

19

problem is the use of one or a very few characters that have been weighted a priori to

structure classifications (see also Garciacutea et al 2009) although it is perhaps not

surprising that such an approach sometimes results in the delineation of taxa that have

indeed turned out to be quite distinct (cf eg Burtt Astragalus versus Oxytropis)

Taxa recognized here are monophyletic yet many are polythetic at the

morphological level that is they lack unique features characterizing (defining) all and

only members of that family they can be recognised phenetically only by the unique

combinations of characters that they posess This is the result of evolution any

synapomorphy characterizing a taxon may be lost or modified beyond easy recognition

in some of its members or the synapomorphy may appear to be identical to a feature

that has evolved in parallel in a quite unrelated plant That plant groups are polythetic is

almost as much a feature of monophyletic taxa as the paraphyletic taxa common in

evolutionary classifications Some families now include substantial variation as

phenetically distinct derived groups are placed in their proper phylogenetic position -

examples are Ericaceae which include the erstwhile Empetraceae a wind-pollinated

group that is florally very different and the overwhelmingly large-flowered and animal-

pollinated Plantaginaceae which now include several derived small-flowered aquatic

and wind-pollinated groups that were previously placed in separate families

Thus some families as delimited here may not be easy to recognize However

remember that detecting relationships - use whatever characters you can even if they

are not obvious - and naming a plant - focus on easy-to-see characters that may not

reflect relationships - are quite different problems Taxa although natural may not be

readily recognizable indeed it was in exactly this context that Lamarck worked out the

basic principles of writing dichotomous keys in 1778 (see Scharf 2007 for keys and the

like) Of course Lamarcks idea of nature was very different from ours - he thought that

there was some kind of continuum of form on which living organisms were to be

situated with no real gaps anywhere - but this meant that his genera (for example)

might well not be sharply distinct from each other even if each were part of the real

continuum that was life Lamarcks separation of the establishment of relationships and

groupings in a system and identification of the plants in that system have been widely

accepted (but cf eg Godfray amp Knapp 2004 many of the contributors to Heywood et

al 2007) Perhaps the best way of identifying plants at the family level is by well-made

multiple access keys as in Watson and Dallwitz (1992a onwards family limits there may

differ substantially from those adopted here) Multiple access keys free users from the

constraints of dichotomous keys in which particular characters are needed at each step

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

20

of the identification process before there can be further progress Instead those

characters that happen to be evident on a specimen can be used in whatever order is

convenient when linked to illustrations glossaries etc their power is enormous (see

Dallwitz et al 2000 [2006] for the principles underlying their construction and use)

Nevertheless dichotomous keys such as those of Hutchinson (1973) and Franz Thonner

(Geesink et al 1981) have their uses Of course taxa coming out adjacent in keys may

well not be at all related

(When identifying large numbers of plants even more efficient than either style of

identification and certainly lots more fun is sight identification Unless you have a

photographic memory you have to build up your knowledge of comparative plant

morphology - on which the ability to make accurate identifications depends - by

repeated observation When faced with an unknown plant I always look for leaf teeth

and stipules distinctive hair types especially stellate and T-shaped hairs glands and

punctations of any sort the presence of latex or other exudate and check leaf insertion

smelling crushed leaves can also be helpful In this context nodal anatomy can usually

be checked using a razor and a hand lens or even simply carefully examining leaf scars

The short paragraphs added after most families may help in confirming familial

identifications)

For the record and for the little that it is worth there are 4 orders and 13 families

of gymnosperms characterised on these pages and together they include some 82

genera and 947 species For angiosperms comparable figures are 56 orders 445

families 13208 genera and 261750 species (of which monocots include 11 orders 89

families 2759 genera and 52760 species) Note however that higher mathematics

was never my strong point and anyway these are pretty meaningless figures even for

species which many (but not all) might concede smacked slightly more of reality than

other taxa estimates range as high as 422000 (Govaerts 2001) Furthermore numbers

of genera and species change daily Nevertheless as emphasized here families are

useful in teaching we as a community can ensure that their limits remain largely stable

and by concentrating on relatively few of them one can gain some familiarity with much

of the worlds flora For summaries of the sizes of orders and families and of the general

arrangement followed in these pages see also the Statistics page itself

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

21

A PHYLOGENETIC TREE OF SEED PLANTS (THE MAIN TREE)

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

22

The Main Tree above is a conservative summary of well supported relationships

between the major clades of extant seed plants Reading - interpreting and

understanding - such a phylogenetic tree is not necessarily easy so reading a tree may

require a bit of practice OHara (eg 1988 1992) provides an accessible introduction to

thinking and talking about history in the context of looking at phylogenetic trees More

recently Krell and Cranston (2004) Crisp and Cook (2005) and others have emphasized

how careful one must be when interpreting and talking about evolution in ladderized

trees in particular and phylogenetic trees in general Gregory (2008) provides a

particularly detailed discussion of this problem In ladderized trees the smaller (in terms

of numbers of terminals) sister taxon is consistently shown on the same side at every

node thus the trees tend to be pectinate like the teeth of a comb however the

horizontal axis that is as a result so evident has no polarity or particular direction The

use of the adjective basal when discussing phylogenetic trees is especially dangerous

(see also Wojciechowski et al 2004 D Soltis et al 2005b) When I use the term and the

context is not otherwise clear I am referring to the pectinations at the base of a

ladderised tree Remember that when talking about sister taxa one can never be basal

to the other or older or younger although members of one can be more derived - in the

sense of having more apomorphies - than the other (but even then one has to be

careful) The words primitive and advanced and lower and higher should

practically never be used especially when talking about taxa Amborellaceae and

Pinaceae are sister to all other flowering plants and Pinales and are basal to the crown

groups of all other flowering plantsPinales respectively but that does not mean they

are thereby primitive Pinaceae in particular have numerous apomorphies The word

plesiomorphic is far less loaded than primitive and can be used to talk about

individual characters

Most trees here have been more or less ladderized as is the one above The

ladderization may be imperfect for example you can see that on the Main Tree

asterids with ten orders follow rosids and relatives which have seventeen although

this is in part because of the currently unresolved nature of relationships between the

main clades of core eudicots (this is likely to change soon) The Main Tree could have

been drawn with Amborella Acorus or a host of other taxa at the far right without

offending any relationships phylogenetic trees are like mobiles the only fixed points

being the nodes Despite the problems just mentioned pectination interpreted

carefully has its value and it is not an arbitrary process As one reads the terminals of a

pectinate tree from left to right adjacent terminals will be separated by apomorphies

that will cumulate in an appropriate fashion When thinking of a book or a herbarium

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

23

sequence (see Haston et al 2007) this is of some value Nymphaeales and

Austrobaileyales are here adjacent on the tree but they could be separated by

hundreds of families in the sequence merely by rotating only the node from which

Nymphaeales and their sister group (all other angiosperms minus about 75 species)

arise If adjacent in a book or herbarium or as in these summary pages then it is

relatively easy to relate their characters both apomorphic and plesiomorphic but if

separated by hundreds of pages or two floors in a large building then it is less easy to

get anything from the sequence (Since all orders in the pages of the main part of this

site are preceded by the apomorphies of all nodes immediately below them in the seed

plant phylogeny and because of the linkages that have been built in to the site this

problem is much less serious there there is no sequence) So the particular sequence of

all seed plants that is based on a phylogenetic classification but is to be used in herbaria

or any other place where taxa have to be arranged linearly as in this part of APweb

will be one that maximizes the number of taxa that are both successive branches of the

tree and placed successively in the sequence Since specimens are generally filed under

families the outline of a new family sequence for arranging herbaria and books can now

be suggested (Haston et al 2007) and such a standardized sequence will further help

teaching and learning about plants

ON THINKING ABOUT APOMORPHIES

Identifying apomorphies is important because understanding the evolution of

morphology in the broad sense - ie including anatomy chemistry etc - in the context

of diversification is one of our major goals For identifying apomorphies several

preconditions must be met One needs to have an accurate well supported phylogeny

one has to have examined the right taxa both from the point of view of morphology and

molecules one has to have coded the characters right (ie delimited states

appropriately) and one has to use an appropriate model of evolution when fitting the

variation to the tree (see eg Omland 1999 Stevens 2006b) If the distributions of

apomorphies below are compared with those in other studies (eg Turgeon et al 2001

Bremer et al 2001 Endress 2001 Albach et al 2001a Judd amp Olmstead 2004 D Soltis

et al 2005b Zhang et al 2006 Judd et al 2007) the reader may find a number of

differences Although I have integrated such studies as far as is possible there are five

reasons that link to the preconditions just mentioned why there may be differences

where features are placed on trees

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

24

1 I may not have found all the information about a particular character there may be

disagreement over its interpretation or I have added information

2 The sampling of nearly all molecular studies is very incomplete (see Salisbury amp Kim

2001 for problems caused by sampling) indeed it is commonplace to decry the

incompleteness of molecular sampling But not only is the sampling in molecular studies

often less than we might wish that of the morphological and chemical characters whose

evolution we are interested in understanding is also often very poor So for many

anatomical chemical and embryological characters that are confidently said to

characterise families and other groups we all too often have no idea if those characters

are applicable to the whole clade or just to a subgroup within that clade However as

our sampling improves we can locate changes on the tree more precisely if in

unexpected positions Thus is is sometimes suggested that Ericaceae have ellagic acid

(Soltis et al 2005b) in fact the little evidence we have indicates that only a small clade

of some 80 species has ellagic acid and the rest of the family - itself alone about one

third of all Ericales - does not Similarly Ericaceae are noted for their

ectendomycorrhizal associations but Enkianthus sister to all other Ericaceae

apparently lacks these (Abe 2005) Thus these mycorrhizae and their associated

distinctive hair roots are unlikely to be apomorphies of Ericaceae - however it appears

that other families near Ericaceae may also have ectendomycorrhizae (see Asai 1934)

3 I am fitting characters to a very conservative tree with many polytomies although the

nodes that are utilised are for the most part strongly supported polytomies make the

optimisation of characters that is the assigment of character state change to a

particular node on the tree notably difficult (eg Madison amp Madison 2002) In nearly

all studies of the evolution of characters distributions of characters are optimised on a

more or less fully resolved tree and the construction of supertrees may yield yet more

detailed hypotheses of relationships (for literature on supertrees see Cotton amp

Wilkinson 2007 2008) Of course some nodes on such fully resolved trees andor

supertrees may have little support and optimisations of characters on such trees may

carry correspondingly little conviction Even parts of some of the trees used here have

poor support eg relationships within aquatic Alismatales etc although I indicate

when this is the case

4 Exactly how one goes about optimising a character on a tree is critically important

Even using simple parsimony optimisations (ACCTRAN or DELTRAN ACCelerated

TRANsitions or DELayed TRANsitions) the position of synapomorphies on trees - and

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

25

hence our ideas of evolution - may differ and this is still more true if one uses maximum

likelihood or Bayesian methods and makes apparently reasonable suggestions about

weighting gains over losses (eg Donoghue amp Ackerley 1996 Cunningham et al 1998

Omland 1997 1999 Ree amp Donoghue 1999 Polly 2001 Webster amp Purvis 2001

Ronquist 2004 Crisp amp Cook 2005) Sannier et al (2007) show how in in Arecaceae that

where on a phylogenetic tree one might peg changes in microsporogenesis will depend

on the methods one uses to do this and Pedersen et al (2007) discuss the sometimes

very substantial effect of node support on the posterior probabilities of ancestral

character states Here I use parsimony optimization not always as explicit as it might be

but I have often indicated where there are particularly important uncertainties as to the

positions of particular character changes on the tree

5 Finally although I have paid quite a lot of attention to the delimitation of the

character states that make up all the characterizations I have not spent enough time on

this critical operation If we are interested in understanding evolution then fitting the

basic variation - not character states - to a tree in principle allows greater flexibility in

understanding morphology in the context of local phylogenies (see also Stevens 2000

Endress 2005c) However many character states used here are delimited globally that

is they are circumscribed in the context of the variation shown by individual characters

across all angiosperms andor in the context of classic ideas of character evolution

Character states often have arbitrary limits and serve best to communicate

information whether they are in fact suitable for either phylogenetic analysis or

understanding evolution are separate issues Studies have rather unsurprisingly

perhaps but importantly shown that dividing the one character into different sets of

states may yield differing ideas of evolution of that character (eg Lamb Frye amp Kron

2003 Hibbett 2004) When looking at trees on which character states are optimised

one should bear in mind the problems surrounding the delimitation of states (eg

Stevens 2000 2006b) and the danger of using pollen or other types - constructs

based on many characters that vary independently but which effectively get lost in

these types thus Blackmore et al (2009) decomposed the pollen types in Asteraceae

into 52 characters

I use Remanes three main criteria of homology or better similarity when

determining the basic similarity of structures on different organisms (see Remane 1952)

These criteria are special properties position and intermediates Special properties

include anatomical or chemical characters gene expression data etc Position refers

to the position of an organ with respect to landmarks on the plant Although plants are

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

26

plants and landmarks are not as fixed as one might like trying to work out the relative

positions of parts is a good way of understanding morphology so long as one does not

try to out-do Procrustes Intermediates include intermediates found on the same

plant or on different plants Intermediates may observed during development or be

adult structures Thus one may find changes during development which suggest what

the nature of a particular structure is and very different stuctures on different plants

may turn out to be similar early in development Or one can simply compare different

taxa and see that structures that appear to be very different when comparing two

different taxa can be linked morphologically by looking at other taxa

Nevertheless the use of these criteria may not yield an unambiguous answer as to

what a structure is even given a solid phylogeny and an improved understanding of

development (see Jaramillo amp Kramer 2007 for a useful discussion) As Endress (2005c)

observed a number of features - position function development shape anatomy

histology gene activity and relationships to other taxa that clearly have petals - can be

used to distinguish a petal (for example) from other floral structures if a petal does not

have one of these features is it thereby not a petal Thus Maturen et al (2005) recently

found that floral organ diversity genes (B and C) were expressed in the large white

inflorescence bracts of Cornus (see also Costa et al 2005) Peney et al (2005) noted that

not all monosulcate pollen grains in monocots have the same developmental pathway

and that as a result such pollen might not have the same ancestral state Reeves and

Olmstead (2003) suggested that the genetic mechanisms causing monosymmetry in

Lamiales and Solanales were different and Serna and Martin (2006) described similar

problems with the development of hairs in Arabidopsis when compared with that of

hairs in Antirrhinum and Solanaceae Indeed as one perhaps might expect delimitation

of states and characters does not necessarily become easier with increasing knowledge

of development etc Thus Buzgo et al (2004) Matthews and Endress (2005) and others

have shown how hard it can be to distinguish between eg prophylls and other floral

structures as their behaviour is studied during the course of the development of the

flower Similarly at what concentration is a particular secondary metabolite deemed to

be present (Waterman (2007)

The validity of the approach used here that of fitting morphological variation to a

largely molecular-based tree may be questioned However I think it rather unlikely that

well-supported molecular branches will be overturned by morphological data Indeed

analyses of morphological data alone do provide support for many of the clades evident

in molecule-only analyses and analyses with morphological and molecular data

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

27

together may lead to increased support for clades (eg Hufford 1992 Nandi et al 1998

[but adding morphological data reduces support for a number of critical clades too]

Doyle amp Endress 2000 however in none of these papers is the use of morphology

without ambiguity) It is unfortunately clear that the use of morphology alone may not

suggest problems in the phylogenetic placement of taxa that later turn out to have been

wrongly included (eg Zhang et al 1992) Thus I have been wary of putting much weight

on clades that have only morphological support but note that I have treated molecule-

based clades with low bootstrap or jacknife support values (esp below 70) or low

posterior probabilies (below 095) likewise Although I may have been mistaken in

placing so much emphasis on molecular data in terms of providing the basic

phylogenetic framework for angiosperms morphological and molecular data are only

very rarely in irreconcilably strong conflict There are indeed a few places where the

conflict seems extreme These include the relative positions of the Monimiaceae and

Hernandiaceae (Laurales) the position of Hanguanaceae (Commelinales [as here] or

Zingiberales) and of Triplostegia (is it in Dipsacaceae or Valerianaceae - see

Dipsacales) Fossils are unlikely to affect the topologies of the trees presented here but

see below for their importance in understanding morphological evolution in general and

the evolution of angiosperms in particular Nevertheless some largely reject the idea

that trees based on molecular data alone can recover phylogenetic relationships

especially when branching points are old and prefer to used trees based on analysis of

morphological data including those taken from fossils (Hilton amp Bateman 2006 Farjon

2007)

All in all however the extent of the congruence between morphological and

molecular data is impressive and heartening and many clades can be characterised

morphologically It seemed in 1998 that there were no unambiguous morphological

synapomorphies for angiosperm orders (K Bremer 2000) and this is still true if by

unambiguous is meant non-homoplasious However many orders and other clades

have synapomorphies even if these may be indistinguishable at least at the current

level of morphological and developmental knowledge from parallel occurences

elsewhere As our morphological knowledge increases so too does the number of

apomorphies

Endress and Matthews (2006a) emphasize the importance of tendencies and

developmental constraints when thinking about characters of clades within the rosids -

many characters or character combinations occur in a rather sporadic fashion within

the clade and apparently notably less frequently outside it There are a number of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

28

examples of tendencies here and in the individual order pages such as the features

enclosed by parentheses in groups above the level of family eg see (cuticular waxes

as aggregated rodlets) for the commelinids and the discussion of the distribution of

polyandry that is flowers with numerous stamens in the asterid I + II groups etc

Tendencies may involve more than single characters When thinking of the

associations of features like integument thickness and vascularization of the integument

(Endress amp Matthews 2006a) one can imagine fairly simple developmental

preconditions being operative As Endress and Matthews (2006a) note it is difficult to

imagine vascular tissue developing in an integument only two cells thick Changes here

would be loosely correlated if morphologically linked However the strongly correlated

changes noted by Givnish et al (2005) are ecologically linked but are presumably

morphologicallydevelopmentally independent When there seem to be characters

evolving more or less together tests can be carried out to see if the changes are

concentrated on certain branches of the tree (eg Maddison 1990 Sanderson 1991

Maddison amp Maddison 2000) Such tests have rarely been carried out at higher levels in

angiosperms

One of the most striking examples of a tendency is the distribution of N-fixation

restricted as it is to a monophyletic group of four clades although it has arisen seven (or

perhaps several more) times independently within the N-fixing clade and several

members of two quite different kinds of bacteria are involved (eg Clawson et al 2004

Elliott et al 2007 Sprent amp James 2007) Other examples of work that bears on the issue

of tendencies include the findings that flowers of polysymmetrical Arabidopsis have

genes like TCP1 that are expressed asymmetrically during early development and TCP1

is a probable orthologue of the well-known CYC gene of Antirrhinum that is involved in

the development of monosymmetric slowers there (Cubas et al 2001 Costa et al 2005

etc) Parallelism might build on this underlying morphologically cryptic monosymmetry

even if details of the genetic mechanisms causing the monosymmetry evident in

particular groups may be different (Reeves amp Olmstead 2003 Cubas 2004 see above)

Similarly there have been several recent suggestions that the capability to synthesise a

particular metabolite may be switched off but not lost and so can sometimes be

reacquired (eg Wink amp Witte 1983 Wink 2003 Liscombe et al 2005 Larsson 2007

Waterman 2007) Hence perhaps the rather spotty distribution of many secondary

metabolites like ellagic acid the indole alkaloid camptothecin iridoids etc when

considered in the context of phylogenies As a non-botanical example - but a rather nice

one - Salwini-Plawen and Mayr (1961) suggested some time ago that there has been

considerable parallelism (40-65 or more independent origins) in the evolution of eyes in

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

29

metazoans However Pax 6 seems to be a master control gene that is involved in eye

formation perhaps throughout all bilateralians (eg Gehring amp Ikeo 1999 Erwin amp

Davidson 2002) which changes the nature of the problem Note however that in

general our invocation of evolutionary tendencies to explain the patterns of

distributions of characters is really an admission that we do not understand those

patterns

As we find out more about variation we will probably find fewer and fewer features

constant throughout a group Most unqualified statements of presence and absence

should properly be qualified as usually present or usually absent if one is thinking of

the characterisations on the order pages as encompassing the total variation within a

clade Thus Pistia alone among monocots as so far known has sieve tube plastids with

starch grains not protein crystals However this does not affect the fact that sieve tube

plastids with cuneate protein crystals and lacking starch grains are an apomorphy for

monocots Furthermore fossils in a number of cases suggest character combinations

unknown in extant taxa as may be seen in the discussions of Fagaceae Platanaceae

Iteaceae Calycanthaceae etc Confusing the issue there may be questions as to where

exactly on the tree a particular fossil is to be placed (eg see Nymphaeaceae

Calycanthaceae Archaefructus etc)

To summarize given our current understandings of both phylogenies and

characters evolution of some characters in which we are interested seems very labile

(see eg D Soltis et al 2005b Endress and Matthews 2006a Stevens 2006b Ekman et

al 2008 for a good recent study) and I have been cautious when talking about character

evolution Much effort must continue to be spent in summarizing characters of well-

established clades at all levels providing features by which they may be recognized and

signaling synapomorphies Remember that (1) the basic morphological anatomical and

chemical knowledge of many critical taxa is woefully incomplete (2) different

assumptions about character evolution may greatly affect the position of

synapomorphies on trees (3) in many cases relationships within and between many

groups are too uncertain at present to worry very much about synapomorphies and

(4) we must be clear about what we do and do not not know As mentioned above I

have indicated in a number of places where there are particularly important

uncertainties as to where characters should be placed on the tree Nevertheless it is a

relatively easy matter to update notes such as these and it can be a simple matter to

incorporate new data on characters that have never before been considered in the

context of a tree Much basic - and unfortunately perhaps unfashionable - work must be

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

30

carried to clarify the distribution of morphological anatomical and chemical characters

for examples of what can to be done see the work of P K Endress and collaborators

(floral morphology and development) and S R Jensen and collaborators (iridoids) But

acquiring information about nodal anatomy cell and tissue distribution nodal anatomy

and the like is not difficult at all and should be generally encouraged

But all this is merely a necesary prelude to the understanding of evolution For this

one needs to know a lot more including dating the phylogeny understanding the

function(s) of characters factoring in relevant aspects of the palaeoenvironment in

which an apomorphy first appeared etc

SUMMARY OF THE SYSTEM

Below is a formalised summary of the relationships within orders of the families of seed

plants There are a few families that are not recognised even as options in APG II and

vice versa as well as a few extra orders The families to be recognised in the most

recent edition of Mabberleys The Plant Book (Mabberley 2008) are also largely

consistent with those below But all differences are trivial and will - I hope - eventually

disappear

Square brackets - [] - enclose clades the plus sign - + - designates sister taxa a

comma - - denotes part of a polytomy and quotation marks - - denotes a

paraphyletic group

SEED PLANTS

GYMNOSPERMS

Cycadales

Cycadaceae + Zamiaceae

Ginkgoales

Ginkgoaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

31

Pinales

Pinaceae [[Araucariaceae [Podocarpaceae + Phyllocladaceae]] [Sciadopityaceae

[Taxaceae + Cupressaceae]]]

Gnetales

Ephedraceae [Gnetaceae + Welwitschiaceae]

ANGIOSPERMSFLOWERING PLANTS

Amborellales

Amborellaceae

Nymphaeales

Hydatellaceae [Cabombaceae + Nymphaeaceae]

Austrobaileyales

Austrobaileyaceae [Schisandraceae + Trimeniaceae]

Chloranthales

Chloranthaceae

MAGNOLIIDS

Magnoliales

Myristicaceae [Magnoliaceae [[Himantandraceae + Degeneriaceae] [Eupomatiaceae +

Annonaceae]]]

Laurales

Calycanthaceae [[Siparunaceae [Gomortegaceae + Atherospermataceae]] [Monimiaceae

[Hernandiaceae + Lauraceae]]]

Canellales

Canellaceae + Winteraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

32

Piperales

[Hydnoraceae + Aristolochiaceae] [Piperaceae + Saururaceae]

MONOCOTS

Acorales

Acoraceae

Alismatales

Araceae [Tofieldiaceae [[Alismataceae [Hydrocharitaceae + Butomaceae]]

[Scheuchzeriaceae [Aponogetonaceae [Juncaginaceae [Maundiaceae [[Posidoniaceae

[Ruppiaceae + Cymodoceaceae]] [Zosteraceae + Potamogetonaceae]]]]]]]

Petrosaviales

Petrosaviaceae

Dioscoreales

Nartheciaceae [[Taccaceae + Thismiaceae] [Burmanniaceae + Dioscoreaceae]]

Pandanales

Velloziaceae Triuridaceae Stemonaceae [Pandanaceae + Cyclanthaceae]

Liliales

Corsiaceae [Campynemataceae [Petermanniaceae [Colchicaceae + Alstroemeriaceae]]

Melanthiaceae [[Philesiaceae + Rhipogonaceae] [Smilacaceae + Liliaceae]]]

Asparagales

Orchidaceae [[Boryaceae [Blandfordiaceae [Lanariaceae [Asteliaceae + Hypoxidaceae]]]]

[[Ixioliriaceae + Tecophilaeaceae] [Doryanthaceae [Iridaceae [Xeronemataceae

[Xanthorrhoeaceae [Alliaceae + Asparagaceae]]]]]]]

COMMELINIDS

Unplaced

Dasypogonaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

33

Arecales

Arecaceae

Poales

[Typhaceae + Bromeliaceae] [Rapateaceae [[Thurniaceae [Juncaceae + Cyperaceae]]

[[Mayacaceae [Eriocaulaceae + Xyridaceae]] [Flagellariaceae [Anarthriaceae

[Centrolepidaceae + Restionaceae]]] [Joinvilleaceae [Ecdeiocoleaceae + Poaceae]]]]]]]

Commelinales

[Commelinaceae + Hanguanaceae] [Philydraceae [Haemodoraceae + Pontederiaceae]]

Zingiberales

Musaceae [Strelitziaceae + Lowiaceae] Heliconiaceae [[Cannaceae + Marantaceae]

[Costaceae + Zingiberaceae]]

Ceratophyllales

Ceratophyllaceae

EUDICOTS

Ranunculales

Eupteleaceae [Papaveraceae [[[Lardizabalaceae + Circaeasteraceae] [Menispermaceae

[Berberidaceae + Ranunculaceae]]]]

Sabiales

Sabiaceae

Proteales

Nelumbonaceae [Platanaceae + Proteaceae]

Trochodendrales

Trochodendraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

34

Buxales

Haptanthaceae [Buxaceae + Didymelaceae]

CORE EUDICOTS

Gunnerales

Gunneraceae + Myrothamnaceae

Dilleniales

Dilleniaceae

Saxifragales

Peridiscaceae [[Paeoniaceae [Altingiaceae [Hamamelidaceae [Cercidiphyllaceae +

Daphniphyllaceae]]]] [[Crassulaceae [Aphanopetalaceae [Tetracarpaeaceae

[Penthoraceae + Haloragaceae]]]] [Iteaceae [Grossulariaceae + Saxifragaceae]]]]

Cynomoriaceae unplaced

Vitales

Vitaceae

ROSIDS

FABIDROSID I

Zygophyllales

Krameriaceae + Zygophyllaceae

Celastrales

Lepidobotryaceae + Celastraceae

Oxalidales

Huaceae [[Connaraceae + Oxalidaceae] [Cunoniaceae [Elaeocarpaceae [Brunelliaceae +

Cephalotaceae]]]]

Malpighiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

35

[Achariaceae [Goupiaceae [Violaceae + Passifloraceae] [Lacistemataceae Salicaceae]]

[Lophopyxidaceae Putranjivaceae] [Ctenolophonaceae [Erythroxylaceae +

Rhizophoraceae]] Linaceae Ixonanthaceae Humiriaceae Irvingiaceae

Centroplacaceae Pandaceae [[Bonnetiaceae + Clusiaceae] [Calophyllaceae

[Hypericaceae + Podostemaceae]]] [Malpighiaceae + Elatinaceae] Ochnaceae

[Peraceae [Rafflesiaceae + Euphorbiaceae]] [Phyllanthaceae + Picrodendraceae]

[Balanopaceae [[Trigoniaceae + Dichapetalaceae] [Chrysobalanaceae +

Euphroniaceae]]] Caryocaraceae

N-FIXING CLADE

Fabales

Quillajaceae [Fabaceae [Polygalaceae + Surianaceae]]

Rosales

Rosaceae [[Barbeyaceae [Elaeagnaceae Dirachmaceae Rhamnaceae]] [Ulmaceae

[Cannabaceae [Moraceae + Urticaceae]]]]

Cucurbitales

Anisophylleaceae [[Corynocarpaceae + Coriariaceae] [Cucurbitaceae [Tetramelaceae

[Datiscaceae + Begoniaceae]]]] Apodanthaceae

Fagales

Nothofagaceae [Fagaceae [[Myricaceae + Juglandaceae] [Casuarinaceae

[Ticodendraceae + Betulaceae]]]]

MALVIDROSID II

pgtGeraniales

Geraniaceae [[Melianthaceae + Francoaceae] [Vivianaceae + Ledocarpaceae]]

Myrtales

Combretaceae [[Onagraceae + Lythraceae] [[Vochysiaceae + Myrtaceae]

[Melastomataceae [Crypteroniaceae [Alzateaceae + Penaeaceae]]]]]

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

36

Crossosomatales

[Staphyleaceae [Guamatelaceae [Crossosomataceae + Stachyuraceae]]] [Aphloiaceae

[Geissolomataceae + Strasburgeriaceae]]

Picramniales

Picramniaceae

Sapindales

Biebersteiniaceae Nitrariaceae [Kirkiaceae [Anacardiaceae + Burseraceae]]

[Sapindaceae [Simaroubaceae Rutaceae Meliaceae]]

Huerteales

Gerradinaceae [Dipentodontaceae + Tapisciaceae]

Malvales

Neuradaceae [Thymelaeaceae [Sphaerosepalaceae Bixaceae [Cistaceae

[Sarcolaenaceae + Dipterocarpaceae]] [Cytinaceae + Muntingiaceae] Malvaceae]]

Brassicales

[Akaniaceae + Tropaeolaceae] [[Moringaceae + Caricaceae] [Setchellanthaceae

[Limnanthaceae [[Koeberliniaceae [Bataceae + Salvadoraceae]] [Emblingiaceae

[Pentadiplandraceae [Gyrostemonaceae Resedaceae Stixis and relatives] Tovariaceae

[Capparaceae [Cleomaceae + Brassicaceae]]]]]]]]

Berberidopsidales

Aextoxicaceae + Berberidopsidaceae

Santalales

Erythropalaceae Olacaceae [[Loranthaceae [Misodendraceae + Schoepfiaceae]]

[Opiliaceae + Santalaceae]] Balanophoraceae unplaced

Caryophyllales

[[Droseraceae [Nepenthaceae [Drosophyllaceae [Ancistrocladaceae +

Dioncophylleaceae]]]]] [[Frankeniaceae + Tamaricaceae] [Polygonaceae +

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

37

Plumbaginaceae]]] [Rhabdodendraceae [Simmondsiaceae [[Asteropeiaceae +

Physenaceae] [[Caryophyllaceae [Achatocarpaceae + Amaranthaceae]]

[Stegnospermataceae [Limeaceae [[Lophocarpaceae [Barbeuiaceae [Aizoaceae

[Sarcobataceae Phytolaccaceae Nyctaginaceae]]]] [Molluginaceae [Halophytaceae

Montiaceae Basellaceae Didiereaceae [Talinaceae [Portulacaceae [Anacampseros etc +

Cactaceae]]]]]]]]]]]]]

ASTERIDS

Cornales

[Cornaceae + Nyssaceae] [Hydrangeaceae + Loasaceae] Hydrostachyaceae [Curtisiaceae

+ Grubbiaceae]

Ericales

[Balsaminaceae [Marcgraviaceae + Tetrameristaceae]] [[Polemoniaceae +

Fouquieraceae] Lecythidaceae [[Sladeniaceae + Pentaphylacaceae] [Sapotaceae

[Ebenaceae [Maesaceae [Theophrastaceae [Myrsinaceae + Primulaceae]]]]]

[Mitrastemonaceae Theaceae [Symplocaceae [Styracaceae + Diapensiaceae]]

[[Sarraceniaceae [Roridulaceae + Actinidiaceae] [Clethraceae [Cyrillaceae +

Ericaceae]]]]]]

LAMIIDASTERID I

Unplaced

Metteniusaceae Oncothecaceae Icacinaceae and some unplaced ex-Icacinaceae

Garryales

Garryaceae + Eucommiaceae

Unplaced

Boraginaceae Vahliaceae

Gentianales

Rubiaceae [Gentianaceae [Loganiaceae [Gelsemiaceae + Apocynaceae]]]

Lamiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

38

Plocospermataceae [Oleaceae [Tetrachondraceae [Calceolariaceae

[Gesneriaceae Peltanthera [Pedaliaceae Carlemanniaceae Martyniaceae Stilbaceae

Plantaginaceae Linderniaceae Bignoniaceae Verbenaceae Lamiaceae Paulowniaceae

Schlegeliaceae Thomandersiaceae Phrymaceae Rehmannia Acanthaceae

Scrophulariaceae Orobanchaceae Byblidaceae Lentibulariaceae]]]]]

Solanales

[Montiniaceae [Sphenocleaceae + Hydroleaceae]] [Convolvulaceae + Solanaceae]

CAMPANULIDASTERID II

Aquifoliales

[[Cardiopteridaceae + Stemonuraceae] [Aquifoliaceae [Helwingiaceae +

Phyllonomaceae]]

Asterales

[Rousseaceae + Campanulaceae] Pentaphragmataceae [[Alseuosmiaceae [Phellinaceae

+ Argophyllaceae]] Stylidiaceae [Menyanthaceae [Goodeniaceae [Calyceraceae +

Asteraceae]]]]

Escalloniales

Escalloniaceae

Bruniales

[Bruniaceae + Columelliaceae]

Apiales

Pennantiaceae [Torricelliaceae [Griseliniaceae [Pittosporaceae [Araliaceae

[Myodocarpaceae + Apiaceae]]]]]

Paracryphiales

Paracryphiaceae

Dipsacales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

39

Adoxaceae [Diervillaceae [Caprifoliaceae [Linnaeaceae [Morinaceae [Dipsacaceae +

Valerianaceae]]]]]

LINKS TO ORDERS AND FAMILIES

[Back to Top]

MAIN GROUPINGS

Asterids commelinids core eudicots asterid 1 asterid

2 eudicots gymnosperms Magnoliophyta monocots N-fixing claderosids seed plants

ALPHABETICAL LISTING OF ALL ORDINAL NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | L | M | N | O | P | Q | R | S | T | U | V | W | X | Z

Abietales Acanthales Acerales Acorales Actinidiales Actinostrobales Adoxales Aescul

ales Agavales Aizoales AkanialesAlismatales Alliales Alseuosmiales Alstroemeriales

Altingiales Amaranthales Amaryllidales Amborellales AmbrosialesAmmiales Amomal

es Ancistrocladales Anisophylleales Annonales Anthobolales Apiales Apocynales Apo

nogetonalesAquifoliales Arales Araliales Aralidiales Araucariales Arecales Aristolochi

ales Athrotaxidales Asarales AsclepiadalesAsparagales Asphodelales Asparagales Ast

eliales Atriplicales Aucubales Austrobaileyales Avenales

Balanitales Balanopales Balanophorales Balsaminales Barbeyales Barclayales Batales

Begoniales BerberidalesBerberidopsidales Betulales Biebersteiniales Bignoniales Bix

ales Boraginales Brassicales Brexiales Bromeliales BrunialesBrunoniales Burmannial

es Burserales Butomales Buxales Byblidales

Cactales Callitrichales Calycanthales Calycerales Campanulales Campynematales Can

ellales Cannales CapparalesCaprifoliales Cardiopteridales Carduales Caricales Carle

manniales Caryophyllales Cassiales Casuarinales CelastralesCentrolepidales Cephalot

ales Cephalotaxales Ceratophyllales Cercidiphyllales Chenopodiales Chironiales Chlor

anthalesChrysobalanales Cinchonales Circaeasterales Cistales Citrales Cocosales Col

chicales Columelliales CombretalesCommelinales Connarales Convolvulales Coriarial

es Cornales Corylales Corynocarpales Crassulales CrossosomatalesCucurbitales Cunn

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

40

inghamiales Cunoniales Cupressales Cycadales Cyclanthales Cymodoceales Cynarales

CynomorialesCyperales Cyrillales Cytinales

Daphnales Daphniphyllales Dasypogonales Datiscales Degeneriales Desfontainiales D

iapensiales Didymelales DillenialesDioales Dioncophyllales Dioscoreales Diospyrales

Dipentodontales Dipsacales Droserales

Ebenales Echiales Elaeagnales Elaeocarpales Elatinales Elodeales Emmotales Empetr

ales Ephedrales Ericales EriocaulalesErythropalaless Escalloniales Eucommiales Euph

orbiales Eupomatiales Eupteleales Euryalaless

Fabales Fagales Falcatifoliales Ficales Flacourtiales Flagellariales Fouquieriales Franc

oales Frangulales

Galiales Garryales Geissolomatales Gentianales Geraniales Gesneriales Ginkgoales G

laucidiales Globulariales GnetalesGoodeniales Greyiales Griseliniales Grossulariales

Grubbiales Gunnerales Gyrocarpales Gyrostemonales

Haemodorales Haloragales Hamamelidales Hanguanales Heisteriales Helleborales He

lwingiales Himantandrales HippuridalesHomaliales Hortensiales Huales Huerteales

Hydatellales Hydnorales Hydrangeales Hydrastidales HydrocharitalesHydropeltidales

Hydrostachyales Hypericales Hypoxidales

Icacinales Illiciales Iridales Irvingales Iteales Ixiales

Jasminales Juglandales Julianiales Juncaginales Juncales

Lacistematales Lactoridales Lamiales Lardizabalales Laurales Lecythidales Ledocarpal

es Leitneriales LentibularialesLigustrales Liliales Limnanthales Linales Liriales Loasal

es Lobeliales Loganiales Lonicerales Loranthales LowialesLythrales

Magnoliales Malpighiales Malvales Marathrales Marcgraviales Mayacales Medusagy

nales Medusandrales MelanthialesMelastomatales Meliales Melianthales Meliosmal

es Menispermales Menyanthales Metteniusales MitrastemonalesMiyoshiales Moni

miales Moringales Musales Myricales Myristicales Myrothamnales Myrsinales Myrta

les

Najadales Nandinales Narcissales Nartheciales Nelumbonales Nepenthales Neuradal

es Nitrariales Nolanales NothofagalesNyctaginales Nymphaeales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

41

Ochnales Oenotherales Olacales Oleales Onagrales Oncothecales Opuntiales Orchid

ales Oxalidales

Saxifragales Pandales Pandanales Papaverales Paracryphiales Parasitaxales Paridales

Parnassiales PassifloralesPenaeales Pennantiales Pentaphragmatales Peridiscales P

etiveriales Petrosaviales Phellinales Philydrales PhyllanthalesPhysenales Phytolaccal

es Picramniales Pinales Pinguiculales Piperales Pittosporales Plantaginales Platanale

sPlumbaginales Poales Podocarpales Podophyllales Podostemales Polemoniales Pol

ygalales Polygonales PontederialesPortulacales Posidoniales Potamogetonales Primu

lales Proteales

Quercales Quillajales Quintiniales

Rafflesiales Ranunculales Rapateales Resedales Restionales Rhabdodendrales Rhamn

ales Rhinanthales RhizophoralesRhodorales Rhoipteleales Roridulales Rosales Rouss

eales Rubiales Ruppiales Rutales

Sabiales Salicales Salvadorales Samolales Samydales Sanguisorbales Santalales Sapin

dales Sapotales SarracenialesSaxegotheales Saxifragales Scheuchzeriales Sciadopitya

les Scleranthales Scrophulariales Scyphostegiales SedalesSilenales Simmondsiales S

milacales Solanales Sphenocleales Sphenostemonales Stangeriales Stellariales Stemo

nalesStilbales Stylidiales Styracales Surianales

Taccales Tamales Tamaricales Taxales Taxodiales Tecophilaeales Terebinthales Tern

stroemiales Theales TheligonalesThymelaeales Tiliales Tofieldiales Torricelliales Tov

ariales Tribelales Trilliales Trimeniales Triuridales TrochodendralesTropaeolales Turn

erales Typhales

Ulmales Urticales

Vacciniales Vahliales Vallisneriales Velloziales Veratrales Verbenales Viburnales Vinc

ales Violales Viscales VitalesVochysiales

Welwitschiales Winterales

Xanthorrhoeales Ximeniales Xyridales

Zamiales Zingiberales Zosterales Zygophyllales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

42

ALPHABETICAL LISTING OF ALL FAMILY NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | K | L | M | N | O | P | Q | R | S | T | U | V | W | X | Y

|Z

Back to Home

In some cases clicking on a family synonym will send you directly to the subfamily in

which the genus of the basionym is to be found

Abietaceae Abolbodaceae Abrophyllaceae Acalyphaceae Acanthaceae [Acanthochla

mydaceae] Acarnaceaee AceraceaeAchariaceae Achatocarpaceae Achradaceae Achy

ranthaceae Acmopylaceae Acoraceae Acristaceae ActaeaceaeActinidiaceae Actinota

ceae Amentotaxaceae Adenogrammaceae Adoxaceae Aegialitidaceae Aegicerataceae

Aegilopaceae Aeginetiaceaee Aegiphilaceae Aesculaceae Aextoxicaceae Agapanthac

eae Agavaceae AgdestidaceaeAgialidaceae Agrimoniaceae Agrostidaceae Ailanthace

ae Aitoniaceae Aizoaceae Akaniaceae Alangiaceae AlchemillaceaeAldrovandaceae A

lismataceae Alliaceae Allioniaceae Allophylaceae Aloaceae Alopecuraceae Alpiniacea

e AlseuosmiaceaeAlsinaceae Alsodeiaceae Alstroemeriaceae Altingiaceae Alzateacea

e Amaranthaceae Amaryllidaceae Amborellaceae Ambrosiaceae Amentotaxaceae A

mmanniaceae Ammiaceae Amomaceae Ampelidopsaceae Amygdalaceae Amyridacea

eAnacardiaceae Anagallidaceae Anarthriaceae Anchusaceae Ancistrocladaceae Andr

omedaceae AndropogonaceaeAndrostachyaceae Androsynaceae Anemarrhenaceae

Anemonaceae Angelicaceae Anisophylleaceae AnnonaceaeAnomochloaceae Anopter

aceae Anrederaceae Anthemidaceae Anthericaceae Anthobolaceae Antidesmataceae

Antirrhinaceae Antoniaceae Aparinaceae Aphanopetalaceae Aphloiaceae Aphyllanth

aceae Apiaceae ApocynaceaeApodanthaceae Aponogetonaceae Aporusaceae Apose

ridaceae Apostasiaceae Aptandraceae AquifoliaceaeAquilariaceae Aquilegiaceae A

raceae Aragoaceae Araliaceae Aralidiaceaee Araucariaceae Arbutaceae Arceuthidace

aeArceuthobiaceae Arctostaphylaceae Arctotidaceae Ardisiaceae Arecaceae Argoph

yllaceae Arisaraceae AristolochiaceaeAristoteliaceae Arjonaceae Armeriaceae Artem

isiaceae Arthrotaxidaceae Artocarpaceae Arundinaceae ArundinellaceaeAsaraceae A

sclepiadaceae Ascyraceaee Asiraceae Asparagaceae Asperulaceae Asphodelaceae As

pidistraceaeeAsteliaceae Asteraceae Asteranthaceae Asteropeiaceae Astragalaceae

Astrocarpaceae AthanasiaceaeAtherospermataceae Atriplicaceae Atropaceae Aucuba

ceae Aurantiaceae Austrobaileyaceae Austrotaxaceae AvenaceaeAverrhoaceae Avet

raceae Avicenniaceae Azaleaceae Azimaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

43

Balanitaceae Balanopaceae Balanophoraceae Balsameaceae Balsaminaceae Bambusa

ceae Barbaceniaceae BarbeuiaceaeBarbeyaceae Barclayaceae Barringtoniaceae Base

llaceae Bataceae Baueraceae Bauhiniaceae Baxteriaceae BegoniaceaeBehniaceae B

elangeraceae Belloniaceae Bembiciaceae Berberidaceae Berberidopsidaceae Berryac

eaee BersamaceaeBertyaceae Berzeliaceae Besleriaceae Betaceae Betulaceae Biebe

rsteiniaceae Bifariaceae Bignoniaceae BischofiaceaeBixaceae Blakeaceae Blandfordia

ceae Blattiaceae Blepharocaryaceae Blitaceae Blyxaceae Boerlagellaceae Bolivaracea

eBombacaceae Bonnetiaceae Bontiaceae Boopidaceae Boraginaceae Borassaceae B

oroniaceae BoryaceaeBotryodendraceae Bougainvilleaceae Boweniaceae Brassicacea

e Brachycaulaceae Bretschneideraceae BrexiaceaeBromeliaceae Brownlowiaceae Br

unelliaceae Bruniaceae Brunoniaceae Brunsvigiaceae Bryoniaceae BuchneraceaeBuci

daceae Buddlejaceae Buglossaceae Bulbocodiaceae Bumeliaceae Bupleuraceae Burc

hardiaceae BurmanniaceaeBurseraceae Butneriaceae Butomaceae Buxaceae Byblida

ceae Byttneriaceae

Cabombaceae Cacaoaceae Cactaceae Caesalpiniaceae Caladiaceae Calamaceae Calce

olariaceae CalectasiaceaeCalendulaceae Callaceae Callicomaceae Calligonaceae Calli

traceae Callitrichaceae Calochortaceae CalophyllaceaeCalthaceae Calycanthaceae Ca

lyceraceae Cambogiaceae Camelliaceae Campanulaceae CampynemataceaeCanacom

yricaceae Canellaceae Cannabaceae Cannaceae Canopodaceae Canotiaceae Cansjera

ceae CapparaceaeCaprariaceae Caprifoliaceae Cardiopteridaceae Carduaceae Carica

ceae Carissaceae Carlemanniaceae CarpinaceaeCarpodetaceae Cartonemataceae Ca

ryocaraceae Caryophyllaceae Casearieaceae Cassiaceae CassipoureaceaeCassythacea

e Castaneaceae Castelaceae Casuarinaceae Catesbaeaceae Cathedraceae Cecropiace

ae CedraceaeCedrelaceae Celastraceae Celosiaceae Celtidaceae Centaureaceae Cen

trolepidaceae Centroplacaceae CepaceaeCephalanthaceae Cephalotaceae Cephalota

xaceae Cerastiaceae Ceratoniaceae Ceratophyllaceae CerberaceaeCercidiphyllaceae

Cercocarpaceae Cercodiaceae Cereaceae Cerinthaceae Ceroxylaceae Cestraceae Cev

alliaceaeChailletiaceae Chamaedoraceae Chamaemoraceae Chamelauciaceae Chaun

ochitonaceae Chelidoniaceae ChelonaceaeChenopodiaceae Chimonanthaceae Chingi

thamnaceae Chionographidaceae Chiranthodendraceae ChironiaceaeChlaenaceae Ch

loanthaceae Chloranthaceae Chloridaceae Chlorogalaceae Chrysobalanaceae Cicerac

eae CichoriaceaeCimicifugaceae Cinchoniaceae Circaeaceae Circaeasteraceae Cissac

eae Cistaceae Citraceae Clematidaceae CleomaceaeClethraceae Clusiaceae Cneorac

eae Cnestidaceae Cnicaceae Cobaeaceae Cochlospermaceae Cocosaceae Coffeaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

44

Colchicaceae Coleogynaceae Columelliaceae Combretaceae Commelinaceae Comocl

adiaceae Compositae CompsoaceaeConantheraceae Connaraceae Conostylidaceae C

onvallariaceae Convolvulaceae Cordiaceae CoreopsidaceaeCoriandraceae Coriariacea

e Coridaceae Corispermaceae Cornaceae Corokiaceae Coronillaceae Correaceae Cor

rigiolaceaeCorsiaceae Corydalaceae Corylaceae Corynocarpaceae Coryphaceae Cost

aceae Cotyledonaceae CoulaceaeCoutariaceae Coutoubiaceae Crassulaceae Crescen

tiaceae Cressaceae Crinaceae Crocaceae CroomiaceaeCrossosomataceae Crotonacea

e Cruciferae Cryptaceae Crypteroniaceae Cryptocorynaceae CryptomeriaceaeCtenol

ophonaceae Cucurbitaceae Cunninghamiaceae Cunoniaceae Cupressaceae Curcumac

eae Curtisiaceae CuscutaceaeCyananthaceae Cyanastraceae Cynanchaceae Cyanella

ceae Cycadaceae Cyclanthaceae CyclantheraceaeCyclocheilaceae Cydoniaceae Cymo

doceaceae Cynaraceae Cynocrambaceae Cynomoriaceae Cyperaceae CyphiaceaeCyp

hocarpaceae Cypripediaceae Cyrillaceae Cyrtandraceae Cyrtanthaceae Cytinaceae

Dacrycarpaceae Dactylanthaceae Damasoniaceae Daphnaceae Daphniphyllaceae Das

ypogonaceae DatiscaceaeDaturaceae Daucaceae Davidiaceae Davidsoniaceae Decais

neaceae Deeringiaceae Degeneriaceae DelphiniaceaeDendrophthoaceae Columelliac

eae Detariaceae Dialypetalanthaceae Dianellaceae Dianthaceae DiapensiaceaeDicha

petalaceae Dichondraceae Diclidantheraceae Dicrastylidiaceae Dictamnaceae Didiere

aceae DidymelaceaeDidymocarpaceae Diegodendraceae Diervillaceae Digitalidaceae

e Dilatridaceae Dilleniaceae Dioaceae DionaeaceaeDioncophyllaceae Dioscoreaceae

Diosmaceae Diospyraceae Dipentodontaceae Diphylleiaceae Diplolaenaceae Dipsaca

ceaeDipterocarpaceae Dirachmaceae Disanthaceae Diselmaceae Dodonaeaceae Do

mbeyaceae Donatiaceae DorsteniaceaeDortmannaceae Doryanthaceae Drabaceae D

racaenaceae Dracontiaceae Drimyidaceae Droseraceae DrosophyllaceaeDryadaceae

Duabangaceae Duckeodendraceae Dulongiaceae Durantaceae Durionaceae Dysphani

aceae

Ebenaceae Eccremidaceae Ecdeiocoleaceae Echiaceae Echinopaceae Ehretiaceae Ela

eagnaceae ElaeocarpaceaeElegiaceae Elatinaceae Ellisiaceae Ellisiophyllaceae Elodea

ceae Elytranthaceae Embeliaceae Emblingiaceae EmmotaceaeEmpetraceae Encepha

lartaceae Engelhardtiaceae Enhalaceae Epacridaceae Ephedraceae Ephemeranthacea

eEpilobiaceae Epimediaceae Eragrostidaceae Eremolepidaceae Eremosynaceae Erica

ceae Erinaceae EriocaulaceaeEriogonaceae Eriospermaceae Erodiaceae Erycibaceae

Eryngiaceae Erysimaceae Erythroniaceae ErythropalaceaeErythrospermaceae Erythro

xylaceae Escalloniaceae Eschscholziaceae Eucomidaceae Eucommiaceae Eucryphiace

aeEuonymaceae Eupatoriaceae Euphorbiaceae Euphrasiaceae Euphroniaceae Eupo

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

45

matiaceae Eupteleaceae EuryalaceaeEustrephaceae Euthemidaceae Eutocaceae Exb

ucklandiaceae Exocarpaceae

Fabaceae Fagaceae Falcatifoliaceae Ferulaceae Festucaceae Ficaceae Ficoidaceae Fit

zroyaceae FlacourtiaceaeFlagellariaceae Flindersiaceae Foetidiaceae Forestieraceae

Fothergillaceae Fouquieriaceae Fragariaceae FrancoaceaeFrangulaceae Frankeniacea

e Fraxinaceae Fraxinellaceae Freycinetiaceae Fritillariaceae Fuchsiaceae Fumariacea

eFunkiaceae

Gaiadendraceae Galacaceae Galanthaceae Galaxiaceae Galedupaceae Galeniaceae G

aliaceae Garciniaceae GardeniaceaeGardneriaceae Garryaceae Geissolomataceae Ge

itonoplesiaceae Gelsemiaceae Geniostomaceae GentianaceaeGeonomataceae Geosir

idaceae Geraniaceae Gerrardinaceae Gesneriaceae Gethyllidaceae Gilliesiaceae Gina

lloaceaeGinkgoaceae Gisekiaceae Gladiolaceae Glaucidiaceae Glechomaceae Glinace

ae Globulariaceae Gnaphaliaceae GnetaceaeGoetzeaceae Gomortegaceae Gomphiac

eae Gomphrenaceae Gonystylaceae Goodeniaceae Gordoniaceae GouaniaceaeGoup

iaceae Gramineae Gratiolaceae Grewiaceae Greyiaceae Grielaceae Griseliniaceae Gr

onoviaceae GrossulariaceaeGrubbiaceae Guaiacanaceae Guamatelaceae Guettardac

eae Gunneraceae Gustaviaceae Guttiferae GyrocarpaceaeGyrostemonaceae

Hachetteaceae Haemanthaceae Haemodoraceae Halesiaceae Hallieraceae Halocarpa

ceae Halophilaceae HalophytaceaeHaloragaceae Hamamelidaceae Hanguanaceae H

aptanthaceae Harmandaceae Hebenstretiaceae HectorellaceaeHederaceae Hedyosm

aceae Hedyotidaceae Hedysaraceae Heisteriaceae Heleniaceae Heliamphoraceae H

elianthaceaeHelianthemaceae Helichrysaceae Heliconiaceae Helicteraceae Heliotropi

aceae Helleboraceae Heloniadaceae HelosidaceaeHelwingiaceae Hemerocallidaceae

Hemimeridaceae Hemiphylacaceae Henriqueziaceae Henslowiaceae Hermanniaceae

Hernandiaceae Herniariaceae Herreriaceae Hesperocallidaceae Heterantheraceae He

teropyxidaceae HeterostylaceaeHibbertiaceae Hibiscaceae Hilleriaceae Himantandra

ceae Hippocastanaceae Hippocrateaceae HippomanaceaeHippophaeaceae Hippurida

ceae Hirtellaceae Holacanthaceae Homaliaceae Hopkinsiaceae Hoplestigmataceae H

ordeaceaeHornschuchiaceae Hortensiaceae Hortoniaceae Hostaceae Houstoniaceae

Huaceae Huerteaceae HugoniaceaeHumbertiaceae Humiriaceae Hyacinthaceae Hyd

atellaceae Hydnoraceae Hydrangeaceae Hydrastidaceae HydrillaceaeHydrocharitacea

e Hydrocotylaceae Hydrogetonaceae Hydroleaceae Hydropeltidaceae Hydrophylacac

eae HydrophyllaceaeHydrostachyaceae Hymenocardiaceae Hypecoaceae Hyperanth

eraceae Hypericaceae Hypopityaceae HypoxidaceaeHypseocharitaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta

ceae ImperatoriaceaeInocarpaceae Inulaceae Ionidiaceae Iriarteaceae Iridaceae Irvi

ngiaceae Isnardiaceae Isophysidaceae IteaceaeStrasburgeriaceae Ixiaceae Ixioliriace

ae Ixonanthaceae

Jalapaceae Jamboliferaceae Japonoliriaceae Jasionaceae Jasminiaceae Johnsoniaceae

Joinvilleaceae JuglandaceaeJulianiaceae Juncaceae Juncaginaceae Juniperaceae Jus

siaeaceae Justiciaceae

Kadsuraceae Kaliphoraceae Kalmiaceae Kaniaceae Kiggelariaceae Kingdoniaceae Kin

giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae

Krameriaceae

Labiatae Lacandoniaceae Lachenaliaceae Lacistemataceae Lactoridaceae Lactucaceae

Lagerstroemiaceaee LamiaceaeLampsanaceae Lanariaceae Langsdorffiaceae Lantan

aceae Lapageriaceae Lardizabalaceae Lasiopetalaceae LauraceaeLawsoniaceae Laxm

anniaceae Lecythidaceae Ledaceae Ledocarpaceae Leeaceae Leguminosae Leitneriac

eae LemnaceaeLennoaceae Lentibulariaceae Lentiscaceae Leoniaceae Leonticaceae

Lepidobotryaceae LepidocarpaceaeLepidocaryaceae Lepidocerataceae Leptaulaceae

Leptospermaceae Lepturaceae Lepuropetalaceae LeuctenbergiaceaeLibocedraceae

Licaniaceae Ligustraceae Lilaeaceae Liliaceae Limeaceae Limnanthaceae Limnocharit

aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace

ae Linderniaceae Lindleyaceae Linnaeaceae LippayaceaeLiriaceae Liriodendraceae Li

ssocarpaceae Littorellaceae Loasaceae Lobeliaceaee Loganiaceae LomandraceaeLoni

ceraceae Lopeziaceae Lophiocarpaceae Lophiolaceae Lophiraceae Lophophytaceae L

ophopyxidaceae LoranthaceaeLotaceae Lowiaceae Lupulaceaee Luxemburgiaceae Lu

zuriagaceae Lychnidaceae Lyciaceae LyginiaceaeLygodisodeaceae Lysimachiaceae Ly

thraceae

Macarisiaceae Mackinlayaceae Madiaceae Maesaceae Magnoliaceae Malaceae Male

sherbiaceae MalortieaceaeMalpighiaceae Malvaceae Mangiaceae Manicariaceae Ma

paniaceae Marantaceae Marathraceae MarcgraviaceaeMartyniaceae Mastixiaceae

Matricariaceae Maundiaceae Mayacaceae Medeolaceae Medusagynaceae Peridiscac

eaeMelaleucaceae Melampyraceae Melanophyllaceae Melanthiaceae Melastomatac

eae Meliaceae Melianthaceae MelicaceaeMeliosmaceae Melittidaceae Melochiacea

e Memecylaceae Mendonciaceae Menispermaceae Menthaceae MenyanthaceaeMe

nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

47

Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

Nypaceae Nyssaceae

Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

ceae Papilionaceae Pappophoraceae PapyraceaeParacryphiaceae Parasitaxaceae Par

ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

48

hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

49

cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

50

ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 6: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

6

conversation If a well-supported hypothesis of monophyly is a necessary prerequisite

for a group to be named it is not a sufficient prerequisite (but cf the PhyloCode -

Cantino amp de Queiroz 2006 Cantino et al 2007) Not all clades need be named indeed

it would barely be practicable (or practical) to do this To decide which clades should be

named additional criteria can be invoked Other things being equal it is helpful if 1 taxa

formally recognised are easily recognizable 2 groups that are well-established in the

literature are preserved 3 the size of groups is taken into account and 4

nomenclatural changes are minimized (Backlund amp Bremer 1998) Thus numerous small

groups have little to recommend them since individually they summarize little

information and tend to clog the memory while groups that are too big may be

amorphous Somewhat similarly Godfray and Knapp (2004 p 562) note that users

want stable informative and accessible classifications that enable easy identification

(see also Simon 2008) - although invoking users without specifying those who make up

this group is not very helpful This classification is for all interested in comparative

biology hence the emphasis on monophyly Although the clades named are sometimes

difficult to characterise there are as we shall see many ways of making such a

classification accessible to all

Problems with this emphasis on monophyly may be caused by reticulation events

such as hybridization endosymbiosis and lateral gene transfer but they are unlikely to

be common confusing factors here Genera can often be pegged to above the level at

which hybridization is at all common However in Poaceae-Pooideae-Triticeae there are

some intractable problems where extremely well established common usage and the

principle of monophyly are likely to remain at odds Many genera are certainly not

monophyletic here being allopolyploids and the genera are ultimately based on the

different genomes they contain (Dewey 1984 Loumlve 1984 Barkworth 2000 for a history

of Triticeae classification Petersen et al 2006) There is also extensive reticulation

reported within Danthonioideae (Pirie et al 2009) Evidence also increases of old

hybridization events elsewhere in flowering plants that at the very least cause

discordance between relationships suggested by different genomic compartments as in

Smedmark and Anderberg (2007 Sapotaceae) and Fehrer et al (2007) Morgan et al

(2009) and Pelser et al (2008 all Asteraceae - hybridization is likely to be quite a

problem at the generic level here) and genera like Medicagoare turning out to have

highly reticulating relationships at the species level (Maureira-Butler et al 2008) Of

course there are many problems trying to apply the concept of monophyly to species

and for many - but not all - biologists strict monophyly is a less important criterion at

this level (Funk amp Omland 2003 see much of the discussion in Houmlrandl 2006)

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

7

The major endosymbiotic events that characterize the clade of which flowering

plants are a part (and gave rise to chloroplasts and mitochondria) are very ancient and

cause no problems for the student of multicellular organisms However lateral gene

transfer has been detected in a number of situations between quite unrelated

organisms (eg Bergthorsson et al 2003 - Amborella and liverworts see also below but

cf Goremykin et al 2009) and it may be particularly common in mitochondria (Sanchez-

Puerta et al 2008) Here too there are no major problems providing one is careful

such transfers do however raise all sorts of interesting biological questions (see

Richardson amp Palmer 2007 for a summary) Although there is increasing evidence for the

importance of genome duplications - hybridization is one cause of this - at various times

during the evolution of seed plants and of palaeopolyploidy events within eg the

Lauraceae and Magnoliaceae clades (Soltis et al 2009 for a summary see eg

the Characters page for further discussion) these too do not currently seem to pose

problems for the adoption of monophyly as the sine qua non of groups to be recognised

formally in this phylogenetic classification but it can make detecting orthologous genes

difficult

The accessory principles of Backlund and Bremer (1998) should be used in

combination Thus keeping the monogenericPlatanaceae separate from its sister

taxon Proteaceae is justifiable Both are much-used names that signal well supported

well defined and easily recognisable groups that have long been recognised as distinct

have several synapomorphies and do indeed look very unlike each other Combining

the two would yield a clade with few obvious apomorphies not to mention the fact

that Nelumbonaceae should by the same logic (it is also monogeneric) also be included

in the expanded family On the other hand it is difficult to justify the continued

recognition of Callitrichaceae or Hippuridaceae monophyletic and distinctive although

they may be If they were recognised several poorly characterised clades would also

have to be carved out ofPlantaginaceae in any classification that aimed to convey a

comprehensive view of the worlds flora The continued recognition

of Valerianaceae and Dipsacaceae also tends to run into this problem (see also Pfeil amp

Crisp 2005 Orthia et al 2005 Albach 2008 etc for useful practical discussions of such

matters) But there are no absolute guidelines If Podostemaceae turn out to be sister

to Hypericaceae (for references here and elsewhere in the Introduction see the

individual families) the subsequent moderate dismemberment of Clusiaceae sl is not

be too high a price to pay for the continued recognition of Podostemaceae Hence the

somewhat provisional recognition of Hypericaceae and Calophyllaceae as well as

Clusiaceae below the families can all be recognized and the name Podostemaceae in

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

8

particular is very well established In a somewhat similar situation if Lemna and its

relatives are a clade sister to most other Araceae should they be recognised as a

separate family Gymnostachys a phenetically fairly distinctive taxon as well as the less

phenetically distinct Orontioideae would have to be recognised as a separate families

(or combined as a single family) too but Araceae in a somewhat restricted sense would

be somewhat more morphologically coherent although not greatly so and not notably

distinct However it is in the very nature of such decisions to be somewhat arbitrary

and unsatisfactory hence the emphasis on consensus classifications here and on

classifications as simply being a means to an end

A useful distinction can be drawn between crown groups and stem groups The

former are monophyletic and include the extant members of a clade and their

immediate common ancestor (see the figure below) The groups characterized in this

site are such groups Thus Proteaceae here are crown group Proteaceae apomorphies

like the single carpel four-merous perianth etc being found in this common ancestor

Stem groups on the other hand include all the members of a lineage immediately after

its split from its sister group and all branches of this lineage In the case of Proteaceae

the stem group would include everything after its split from its sister group

Platanaceae Obviously most of the organisms in the lineage that terminates in crown-

group Proteaceae are unknown only a few being known as fossils and it is not known

exactly where particular apomorphies of crown group Proteaceae evolved along this

lineage

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

9

Indeed there is no reason other than convention or convenience why any group

should not be segregated into several smaller monophyletic groups or merged to

produce a larger unit we can talk about one large thing or about several smaller things

Thinking about aspects of size findings in ethnobiology and cognitive psychology can be

used to suggest that a moderate number - probably fewer than 500 - of families is a

reasonable goal at which to aim and that groupings of taxa throughout any system

should be rather small in size (eg Berlin 1992 Stevens 1994 1997) Major systems such

as those of Linnaeus and Bentham and Hooker were constructed explicitly so as to ease

the burden on the memory (Stevens 1997 2002 see also Scharf 2007) the latter in

particular ensuring that all groups in their classification were relatively small often

containing three to eight immediately subordinate taxa - but by no means all their

groups were formally named Along the same lines Burtt (1977b) suggested that the

number of names at any rank should be at most one third those at the immediately

lower rank - and monotypic taxa might not need a formal name Consistent with such

ideas a fairly broad view of families and orders is taken here whenever the constraints

of monophyly and other criteria used when constructing classifications (see above)

permit The APG system is thus a convention consistent with ideas such as those

expressed by Backlund and Bremer (1998) for similar conventions see eg the Grass

Phylogeny Working Group (2001) and for fungi Hibbett et al (2007)

There are other views Thus Takhtajan (1997) has suggested that smaller families

are more natural This is incorrect Monophyletic groups that include fewer taxa -

Takhtajans smaller families - do not necessarily have more apomorphies than larger

groups even if members of smaller groups will always have more features in general in

common than the larger group that includes them That is they will have their

apomorphies their unique features as well as progressively more plesiomorphies

features found both in the small groups and in the larger clades of which they are a part

having more features in common is indeed one common meaning of more natural

(Note that the implication of the word natural has long been a group of the kind

[usually unspecified] that I think should be recognised and if other botanists had

different ideas too bad their groups were necessarily not natural [see Bather 1927] It

is thus rarely a helpful word and is not used here) Furthermore if this approach is

adopted we will find a slippery slope ahead By this kind of argument all families should

be very small since their members will have a great deal in common and so will be

most natural However as families (for example) are split the relationships that are

evident between the segregates and that were responsible for their being placed in a

single family in the first place will seem to necessitate the recognition of a new order

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

10

etc as is evident in Takhtajans own work - general taxonomic inflation is the result (see

also comparable suggestions in a cladistic context for Brassicales in particular - Ronse de

Craene amp Haston 2006) Such splitting is also questionable when teaching and learning

families since the student needs to understand the system as a whole However for

some genera removed from the families that until now have included them the

phenetic-classificatory-phylogenetic structure in their new home may mandate the

recognition of small families On the other hand Takhtajans suggestion that narrowly

defined families are more useful for phylogenetic studies may be true Indeed I have

more than once regretted prematurely combining groups whether species (in the

context of monographic work) or families (in the course of preparing these notes) but

this is largely a separate issue

Van Steenis (1978) Philipson (1987b) and others have rightly questioned the value

of splitting a group when ideas of the relationships of its constituent members have not

changed - that is very good reasons have to be provided for splitting a family if the

genera within it remain part of the same clade rather than belonging to another clade

Thus APG (2003) broadened the circumscription of Malvaceae because of the

parapolyphyly of some of the families that had historically been associated with it (Judd

amp Manchester 1997 Alverson et al 1999 Bayer et al 1999) These families particularly

Tiliaceae and Sterculiaceae were not at all easy to distinguish their close relationship

(see eg Brown 1814) and overall similarity had long been conceded and to some

workers at least their combination has come as something of a relief Although most of

the larger clades within Malvaceae sl remain difficult to distinguish even with flowers

Cheek (2007) opts for a wholesale and novel dismemberment into ten families

however the very good reasons for doing this are wanting

The same principles are of course applicable when it comes to dividing genera little

other than a headache is gained by splitting genera such

as Drosera and Gnetum (Doweld 2000) as has recently been proposed Thus if an

established genus divides into two (or more) clades this is not a signal for recognising

two groups at the same level - so here it could be argued that the dismemberment

of Pterostylis (Jones amp Clements 2002b) was somewhat unfortunate Along the same

lines if a newly-discovered taxon is sister to an existing named taxon - say a genus - this

does not necessitate the description of a separate genus for the newly described species

(cf Davis 2002) (Of course some [eg Thorne 1976] have suggested that the sizes of

gaps between groups at the same rank should be similar but any principle like this is

inherently flawed since morphological gaps are more unstable than phylogenetic

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

11

relationships and applying it across all flowering plants would both be difficult and cause

substantial changes - see also below) Humphreys and Linder (2009) provide a well-

documented survey of generic concepts in plants which the reader should consult they

note that generic limits (broad versus narrow) have oscillated historically and that

currently larger genera tend to be recognised because studies tend to be on a broader

scale than in the past

Note that invoking similarity or difference - whether qualified (considerable

similarities substantial differences) or not - in a cladistic context as justification for

combining or splitting taxa is not a particularly strong argument (see eg

CardiopteridaceaeStemonuraceae - Karingrehed 2002c) Similarity and difference can

neither be defined precisely since what may seem to be substantial similarities to me

may not to the next person nor are they likely to be stable in the face of our changing

knowledge of morphology and what might be synapomorphies

I might have prefered to merge some families recognised here or split others but

by and large I do not think my own preferences matter very much - and I take the same

position with regards to comparable preferences expressed by others Indeed the

bottom line is that in flagged hierarchies of the kind used here the limits of any

monophyletic unit generally taught and discussed particularly other than species can

be established only by convention and consensus (eg Stevens 2002 2006a Entwisle amp

Weston 2005) This is rather different from the reasonable observation that the

phylogeny itself cannot be achieved by consensus (Thorne 1976) for in most cases there

is a fact of the matter when it comes to relationships Given the increasing support for

the outlines of angiosperm phylogeny a stable consensus classification based on this

phylogeny seems attainable Indeed in addition to providing current ideas of

relationships of seed plants in a synthesised form this site is part of an attempt to build

such a consensus about the circumscription of taxa (see APG 1999 2003 2009 Grass

Phylogeny Working Group 2001 Mabberley 2008) Reaching such a consensus is vital

since what we know of angiosperm phylogeny allows a very large number of

classifications to be based on it and as we find out more the number of possible

classifications increases greatly Unfortunately however nature does not dictate what

the classification should be All classifications are constructed by humans to

communicate particular aspects of groups and relationships Our goals as systematists

are surely to produce robust hypotheses of relationships to understand the evolution of

morphology and the like - but not to argue ad nauseam whether something should be

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

12

a family or a subfamily That way surely lies madness and worse the discredit of our

discipline

There are similar issues whatever naming system is used Thus in phylogenetic

naming (Baum et al 1998 for an example but cf Baum et al 2004 for the PhyloCode

see Cantino amp De Queiroz 2006) an unflagged hierarchy is used in which any

terminations of names used are uninformative about the relative position of taxa If one

adopts the principle of phylogenetic naming one indeed does not have to worry about

the nomenclatural consequences caused by lumping or splitting any well-supported

clade can be named without affecting the name of more or less inclusive clades

Unfortunately unflagged hierarchies have very serious deficiences as communication

devices because they lack one aspect essential in language biological or otherwise -

they contain no intrinsic information about the relationships of the group in question to

others (eg Pfeil amp Crisp 2005 Stevens 2006a) Recent suggestions for using prefixes like

Apo- and Pan- to PhyloCode names will however allow limited information of this

kind to be conveyed but only as it pertains to individual branches and current

proposals do not even mandate that the prefixes be employed consistently In any

event such proposals simply prevent the potential tripling of the number of quite

different names used to describe different aspects of a phylogenetic tree over those

used to name monophyletic groups pure and simple In general where n is the number

of extant species in a group the number of clades in such a group = n-1 (Species will

also need names too for their names see Dayrat et al 2008) Importantly here too

consensus over the clade names commonly learned by students and used in herbaria is

needed otherwise communication will be impeded the names themselves will provide

no guidelines as to which should be chosen The situation is of course more complicated

than this Terminations that convey ideas of rank in a phylogenetic classification can also

be used in phylocode names - however there they will carry no implications of rank

How they will be used is another matter of course

Of course there are other philosophies of classification and some still prefer

evolutionary classifications There classificatory principles differ substantially from those

followed here eg the recognition of paraphyletic taxa may be permitted however

detailed reasons for prefering the taxa that are recognised are rarely given although

nature and natural groups are often mentioned (cf Stuessy amp Koumlnig 2008) For

summaries of commonly used systems see Brummitt (1992) and Mabberley (2008)

new evolutionary systems appear every year or so However even those who allow or

promote the recognition of paraphyletic groups (eg Grant 2003 Thorne 2007

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

13

Heywood et al 2007) may find it of interest to examine a system recognizing only

monophyletic groups it provides a rather different understanding of evolution

Evolutionary classifications in general try and combine phylogeny and

morphological gaps although that is no easy thing to do - it is akin to combining chalk

and cheese (for an attempt to make this impossible task seem to be more objective see

Stuessy amp Koumlnig 2008) However there is a principle from evolutionary classification

that is relevant and I think quite useful here The size of the gap between two groups

tends to be inversely proportional to the sizes of the groups involved (Davis amp Heywood

1963) One can imagine a situation in which a large group is formally divided even

although the distinguishing characters of the two are weak whereas a smaller group

similarly divisable will be left intact

To summarize If hypotheses of phylogeny remain stable we should be able to base

a stable classification on that phylogeny and then get on with our work that is testing

the phylogenies we have elucidating phylogenies in areas where relationships are

unclear studying the evolution of morphology describing species etc In this context

the spread of the Angiosperm Phylogeny Group system (see below) and its widespread

utilisation in technical literature also floras (eg van der Meijden 2005) textbooks (eg

Simpson 2006 Judd et al 2007 [third edition]) dictionaries (Mabberley 2008) more

popular literature (eg Souza amp Lorenzi 2005 Spears 2006) and as an outline for a new

herbarium sequence (Haston et al 2007) is gratifying The posibility that one might be

able to develop a stable phylogeny-based classification of families and in particular

orders represents a dramatic turn-around from the pessimistic attitude about such

higher-level groupings expressed by Davis and Heywood (1963) and Thorne (1976) the

latter even suggesting that we should bury forever the metaphor of the phylogenetic

tree as highly unrealistic (ibid p 56) Returning to Godfray and Knapps (2004) users

of classifications who want a stable informative and accessible classification that

enables easy identification - unfortunately they want cake with everything and cannot

get it - these pages attempt to satisfy as many of their needs as possible but phylogeny

and monophyly are the primary shapers of the APG classification

On this classification in particular

Here I very largely follow the Angiosperm Phylogeny Group classification (APG

2003) Any differences are not to be interpreted as differences in principle simply that

new phylogenies continue to be published and here I attempt to provide an overview

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

14

of current ideas of higher-level relationships of seed plants The Angiosperm Phylogeny

Group classification is based on well-supported relationships evident in the numerous

molecular studies that began to appear in the late 1980s (see APG 1999 for the

principles underlying the classification) but recent phylogenetic work does not

contradict the major outlines of the trees used by APG II (2003) or even those in APG

I (APG 1999) The APG classification has for the most part been conservative and I

am similarly conservative here For the names and authorities I follow APG III (2009)

although the names of the authors may not always be correct these being in something

of a state of flux for the superordinal names used see Chase and Reveal (2009)

although classification at this level here is not exhaustive - only clades with several

orders have superordinal names

Some changes in our ideas of relationships and hence in the clades we talk about

are particularly likely in parts of Lamiales Caryophyllales and Malpighiales for example

although this will not affect the orders themselves Thus some changes to clade

circumscriptions are to be expected although I do not expect them to be substantial

but changes are neither a defect of cladistics nor a necessary consequence of the use of

molecular data Clades are hypotheses of relationships and as hypotheses they may be

overturned However work since APG I has not suggested other than minor changes

in the compositions of the orders even if the odd genus or even family is turning out to

be seriously misplaced - recent examples are Hydatellaceae (from monocots-Poales to

Nymphaeales Saarela et al 2007) Guamatelaceae (from Rosales-Rosaceae to

Crossosomatales Oh amp Potter 2006) and Perrottetia and Bhesa (from Celastrales-

Celastraceae to Huerteales and Malpighiales respectively Zhang amp Simmons 2006) The

main changes have been clarification of the relationhips of individual families or groups

of families that were of uncertain position eg of Chloranthaceae (Moore et al 2007)

Ceratophyllaceae (Jansen et al 2007) and unplaced asterid II families (Winkworth et al

2008a) As mentioned above phylogenies in the area of main interest in these pages are

overwhelmingly tree-like Although there is evidence for acquisition of host plant genes

by parasites (see below) even wider but inexplicable transfer of mitochochondrial

genes in plants like Amborella (Bergthorsson et al 2004 cf Goremykin et al 2009a)

and even transfers of nuclear genes (Vallenback et al 2008) these are the exception

rather than the rule

In cases where the Angiosperm Phylogeny Group II allowed alternatives as to the

limits of families - Papaveraceae in the broad sense or Papaveraceae plus

Pteridophyllaceae plus Fumariaceae Proteaceae in the broad sense or Proteaceae plus

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

15

Platanaceae - the choices made here follow common usage eg as in textbooks like

Judd et al (2007) and Simpson (2006) and particularly in the new edition of

Mabberleys The Plant Book (Mabberley 2008) This latter is itself an attempt to reflect a

consensus the result of taking the opinions of botanists at several meetings A largely

similar consensus is reflected in the APG III classification (APG 2009) where you will

see that alternative classifications have been dispensed with For many people the

existence of such alternative classifications will simply confuse so agreement over

which groupings to use when alternatives are permitted and then dispensing with the

alternatives that are not commonly used seems reasonable I am making the few

changes that are needed to bring the classification here in line with these others but

only gradually since there is not enough time in each day

As already mentioned higher-level relationships in general and the composition of

orders in particular have always presented something of a challenge to systematists

(eg Davis amp Heywood 1963) That being said the composition of some clades like

Apiales Crossosomatales and Pandanales is decidedly unexpected However it is

interesting to see that these higher level clades are generally accepted even in works

with different classificatory philosophies (eg Heywood et al 2007) For clades like

Malpighiales many of the family groupings within Asparagales etc attempts to find

distinctive characters have largely failed (but see Endress amp Matthews 2006a also

the Apomorphies page here) Interestingly as with families some groupings suggested

by molecular studies are supported by morphological andor chemical characters that

have long been known sometimes for over a hundred years the relationship between

Pittosporaceae and ApiaceaeAraliaceae are a case in point (Hegnauer 1969b and

references) As our knowledge of morphology and chemistry improves we can hope for

improvements in the characterisations of clades at all levels

In many cases the new family limits of the Angiosperm Phylogeny Group (see

APG 1999 2003) are not really controversial although changes from the limits

commonly accepted only a decade ago are sometimes quite dramatic (eg Wagenitz

1997) Thus the split of the old Saxifragaceae sl is necessitated by its extreme

polyphyly as also with Icacinaceaesl and Cornaceae sl However the limits and

relationships of such groups had long been considered to be unsatisfactory but until

recently there had been no real reason to prefer one proposed arrangement over

another It is generally accepted that the limits of Lamiaceae and Verbenaceae have to

be redrawn although there is as yet no compelling evidence that the redrawn taxa are

not sister taxa (there is no evidence that they are) But whatever their relationships the

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

16

content of the clades has changed considerably and incidentally they are now easier to

identify than before the decision to recognise two families is not difficult The same is

true for Salicaceae and Achariaceae (Malpighiales) two previously small families that

have received the bulk of the old Flacourtiaceae Clade and hence taxon limits remain

difficult around Theaceae (Ericalessee alsoSladeniaceae Pentaphylacaceae)

and Euphorbiaceae (Malpighiales see

also Phyllanthaceae Picrodendraceae Putranjivaceae etc) Here current groupings

may not represent quite such dramatic changes in our understanding of relationships

Indeed parts of the old Euphorbiaceae that were separated may yet go back together

particularly Phyllanthaceae and Picrodendraceae (see Wurdack et al 2004 Davis et al

2005) even if Theaceae should indeed be dismembered (Geuten et al 2004) However

given that there is no molecular evidence that currently warrants combining all the

segregates of Euphorbiaceae and even if some do come together the clades the

families represent suggest novel groupings not recognised in current classifications

Furthermore Rafflesiaceae appear to be embedded within Euphorbiaceae s str (Davis

et al 2007) so the family is divided Of course maintaining even a moderately narrowly-

circumscribed Euphorbiaceae would entail reducing the iconic Rafflesiaceae to

synonymy Relationships in core Caryophyllales especially

around Phytolaccaceae andNyctaginaceae and also Portulacaceae are incompletely

understood and refashioning of taxon limits will doubtless be needed as cladistic

relationships become apparent (eg see Nyffeler 2007 Ogburn 2007a b) Some

groupings in the old Icacinaceaeand Olacaceae also remain unclear

The discovery of the relationships of parasitic and aquatic groups have presented a

particular challenge to systematists Morphologically some of these plants are so highly

modified that interpretation of the plant body in conventional terms is difficult or even

impossible Thus parasitic groups (broad-leaved angiosperms only) such as Rafflesiaceae

are hard to place since both the vegetative body and the flowers are changed almost

beyond recognition (flowers of rafflesiaceae are a good example of gigantism - Davis et

al 2007) Furthermore plastid gene sequences may be difficult or impossible to obtain

the chloroplast DNA in particular being highly degraded and the rate of molecular

change in general may be high so the problem of long-branch attraction is serious (eg

Duff amp Nickrent 1997 Nickrent et al 1998 Caddick et al 2002a G Petersen et al

2006b) Echlorophyllous sapromycoheterotrophic taxa mostly known from monocots

present similar problems However progress is being made placements for

Apodanthaceae Rafflesiaceae Mitrastemonaceae Cytinaceae and Cynomoriaceae have

recently been suggested (Barkman et al 2004 2007 Davis amp Wurdack 2004 Nickrent et

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

17

al 2004 Davis et al 2007) it seems likely that Burmanniaceae sl are polyphyletic but

both parts are in Dioscoreales (Merckx et al 2006) and relationships within the largely

hemiparasitic Santalales are also gradually being clarified (Maleacutecot 2002) The inclusion

of parasitic taxa in general molecular analyses can cause conniptions (eg Nickrent et al

2004 Davis et al 2004 Chase et al 2006 G Petersen et al 2006b) and there can be

horizontal transmission of genes (eg Davis amp Wurdack 2005 Vitaceae to Rafflesiaceae

Barkman et al 2007 the mitochondrial atp1 gene commonly moves)

Similarly in water plants neither vessels in particular nor much xylem in general is

needed leaves are highly modified and water-mediated pollination if adopted may

well be associated with major changes in floral morphology Here too recent molecular

studies suggest that aquatic groups with hitherto problematic relationships may find

homes Thus Podostemaceae are close to Clusiaceae and Calophyllaceae and are sister

to Hypericaceae (Malpighiales Kita amp Kato 2001) Hydatellaceae which used to be in

Poales are part of Nymphaeales (Saarela et al 2007) and Hydrostachyaceae may be

close to Hydrangeaceae (Cornales Xiang et al 2002) or perhaps should be placed in

Lamiales note that in the first two cases in particular there are morphological and

chemical features that support such a move If Podostemaceae are indeed close to

Clusiaceae I look forward to seeing hypotheses to explain how the dramatic changes in

the vegetative body that have made Podostemaceae so problematic for generations of

systematists took place That conventional wisdom has trouble in understanding or

explaining how the morphologies of groups like Clusiaceae and Podostemaceae can be

related is largely a problem with conventional wisdom Ceratophyllaceae seem to be

finding a position as sister to eudicots (see Moore et al 2007) but their morphology is

so derived that there is no morphological evidence of which I am aware for this

relationship There are also similar although less extreme morphological problems with

many plants pollinated by wind

In such situations especially with some water plants mycoheterotrophs and

parasites a variety of characters may be affected and particularly under such

circumstances groups have tended to be formed using a few or even only a single

character that seems to provide evidence of relationships Add to this the tendency to

weight some characters particularly strongly on a priori grounds and the result was

either the recognition of conglomerate taxa such as Amentiferae which now appear to

be highly polyphyletic or the segregation of families like Plantaginaceae s str (now

much expanded) and Leitneriaceae (now in Simaroubaceae) - all of these are more or

less wind pollinated

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

18

Even although relationships of many parasitic and aquatic groups are being

clarified and at least some of the proposed phylogenetic positions seem to be well

supported it is indeed often still very difficult to understand how such groups have

evolved from their more morphologically conventional relatives However we have had

all sorts of preconceptions as to how characters should evolve Thus we used to

assume that features like highly scalariform vessel perforation plates or the complete

absence of vessels or a flower with an androecium that had many stamens a superior

ovary or separate petals were necessarily plesiomorphic or primitive and conversely

a simple perforation plate and the presence of vessels an androecium with few

stamens inferior ovary or petals that were connnate were almost necessarily

apomorphic or advanced (but cf eg Stebbins 1951) Such assumptions are incorrect

(eg Soltis et al 2005b) Carpels may become secondarily free carpels may fail to close

the seeds then developing outside the confines of the carpel as in some Aspagaraceae-

Nolinoideae Violaceae Berberidaceae Malvaceae-Sterculioideae etc) in Peliosanthes

teta perhaps the only species in Peliosanthes(Asparagaceae-Nolinoideae) the ovary

varies from superior to inferior (Jessop 1976 species limits here need close

investigation see also Kuzoff et al 2001 and Soltis amp Hufford 2002 Saxifragaceae

Apiales etc) many-seeded carpels can evolve from few-seeded carpels

(Razafimandimbison et al 2008) monoecy may be derived from dioecy (Schaefer amp

Renner 2010 and references) Classic studies such as those by Babcock (eg 1947)

on Crepis that assumed that evolution - in this case of the karyotype in particular - was

unidirectional have needed comprehensive re-evaluation (Enke amp Gemeinholzer 2008)

Most if not all characters have reversed andor evolved in parallel as is clear in the

discussion of ovary position here especially in Poales Saxifragales and Asparagales and

also in comments on such morphologically distinctive taxa (when compared with their

immediate relatives) such as Menyanthaceae and especially Pittosporaceae Parallel

evolution may occur even at the level of amino acid substitution as in the independent

acquisition of the phosphoenolpyruvate carboxylase (pepC) gene in C4 photosynthesis in

grasses (Christin et al 2007b see also Blaumlsing et al 2000)

Thinking of how such characters have been used in the past in classifications at the

generic level the results have been similar to those just discussed above for parasitic

plants and others Relying too much on animal pollination syndromes to mark generic

boundaries has all too often led to taxa that are highly unsatisfactory phylogenetically

(see eg Acanthaceae Bignoniaceae Campanulaceae Ericaceae Lamiaceae

Orchidaceae) and over-reliance on characters of fruit and seed (see eg Brassicaceae

and Apiaceae) has also led to unsatisfactory generic limits Again the more general

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

19

problem is the use of one or a very few characters that have been weighted a priori to

structure classifications (see also Garciacutea et al 2009) although it is perhaps not

surprising that such an approach sometimes results in the delineation of taxa that have

indeed turned out to be quite distinct (cf eg Burtt Astragalus versus Oxytropis)

Taxa recognized here are monophyletic yet many are polythetic at the

morphological level that is they lack unique features characterizing (defining) all and

only members of that family they can be recognised phenetically only by the unique

combinations of characters that they posess This is the result of evolution any

synapomorphy characterizing a taxon may be lost or modified beyond easy recognition

in some of its members or the synapomorphy may appear to be identical to a feature

that has evolved in parallel in a quite unrelated plant That plant groups are polythetic is

almost as much a feature of monophyletic taxa as the paraphyletic taxa common in

evolutionary classifications Some families now include substantial variation as

phenetically distinct derived groups are placed in their proper phylogenetic position -

examples are Ericaceae which include the erstwhile Empetraceae a wind-pollinated

group that is florally very different and the overwhelmingly large-flowered and animal-

pollinated Plantaginaceae which now include several derived small-flowered aquatic

and wind-pollinated groups that were previously placed in separate families

Thus some families as delimited here may not be easy to recognize However

remember that detecting relationships - use whatever characters you can even if they

are not obvious - and naming a plant - focus on easy-to-see characters that may not

reflect relationships - are quite different problems Taxa although natural may not be

readily recognizable indeed it was in exactly this context that Lamarck worked out the

basic principles of writing dichotomous keys in 1778 (see Scharf 2007 for keys and the

like) Of course Lamarcks idea of nature was very different from ours - he thought that

there was some kind of continuum of form on which living organisms were to be

situated with no real gaps anywhere - but this meant that his genera (for example)

might well not be sharply distinct from each other even if each were part of the real

continuum that was life Lamarcks separation of the establishment of relationships and

groupings in a system and identification of the plants in that system have been widely

accepted (but cf eg Godfray amp Knapp 2004 many of the contributors to Heywood et

al 2007) Perhaps the best way of identifying plants at the family level is by well-made

multiple access keys as in Watson and Dallwitz (1992a onwards family limits there may

differ substantially from those adopted here) Multiple access keys free users from the

constraints of dichotomous keys in which particular characters are needed at each step

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

20

of the identification process before there can be further progress Instead those

characters that happen to be evident on a specimen can be used in whatever order is

convenient when linked to illustrations glossaries etc their power is enormous (see

Dallwitz et al 2000 [2006] for the principles underlying their construction and use)

Nevertheless dichotomous keys such as those of Hutchinson (1973) and Franz Thonner

(Geesink et al 1981) have their uses Of course taxa coming out adjacent in keys may

well not be at all related

(When identifying large numbers of plants even more efficient than either style of

identification and certainly lots more fun is sight identification Unless you have a

photographic memory you have to build up your knowledge of comparative plant

morphology - on which the ability to make accurate identifications depends - by

repeated observation When faced with an unknown plant I always look for leaf teeth

and stipules distinctive hair types especially stellate and T-shaped hairs glands and

punctations of any sort the presence of latex or other exudate and check leaf insertion

smelling crushed leaves can also be helpful In this context nodal anatomy can usually

be checked using a razor and a hand lens or even simply carefully examining leaf scars

The short paragraphs added after most families may help in confirming familial

identifications)

For the record and for the little that it is worth there are 4 orders and 13 families

of gymnosperms characterised on these pages and together they include some 82

genera and 947 species For angiosperms comparable figures are 56 orders 445

families 13208 genera and 261750 species (of which monocots include 11 orders 89

families 2759 genera and 52760 species) Note however that higher mathematics

was never my strong point and anyway these are pretty meaningless figures even for

species which many (but not all) might concede smacked slightly more of reality than

other taxa estimates range as high as 422000 (Govaerts 2001) Furthermore numbers

of genera and species change daily Nevertheless as emphasized here families are

useful in teaching we as a community can ensure that their limits remain largely stable

and by concentrating on relatively few of them one can gain some familiarity with much

of the worlds flora For summaries of the sizes of orders and families and of the general

arrangement followed in these pages see also the Statistics page itself

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

21

A PHYLOGENETIC TREE OF SEED PLANTS (THE MAIN TREE)

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

22

The Main Tree above is a conservative summary of well supported relationships

between the major clades of extant seed plants Reading - interpreting and

understanding - such a phylogenetic tree is not necessarily easy so reading a tree may

require a bit of practice OHara (eg 1988 1992) provides an accessible introduction to

thinking and talking about history in the context of looking at phylogenetic trees More

recently Krell and Cranston (2004) Crisp and Cook (2005) and others have emphasized

how careful one must be when interpreting and talking about evolution in ladderized

trees in particular and phylogenetic trees in general Gregory (2008) provides a

particularly detailed discussion of this problem In ladderized trees the smaller (in terms

of numbers of terminals) sister taxon is consistently shown on the same side at every

node thus the trees tend to be pectinate like the teeth of a comb however the

horizontal axis that is as a result so evident has no polarity or particular direction The

use of the adjective basal when discussing phylogenetic trees is especially dangerous

(see also Wojciechowski et al 2004 D Soltis et al 2005b) When I use the term and the

context is not otherwise clear I am referring to the pectinations at the base of a

ladderised tree Remember that when talking about sister taxa one can never be basal

to the other or older or younger although members of one can be more derived - in the

sense of having more apomorphies - than the other (but even then one has to be

careful) The words primitive and advanced and lower and higher should

practically never be used especially when talking about taxa Amborellaceae and

Pinaceae are sister to all other flowering plants and Pinales and are basal to the crown

groups of all other flowering plantsPinales respectively but that does not mean they

are thereby primitive Pinaceae in particular have numerous apomorphies The word

plesiomorphic is far less loaded than primitive and can be used to talk about

individual characters

Most trees here have been more or less ladderized as is the one above The

ladderization may be imperfect for example you can see that on the Main Tree

asterids with ten orders follow rosids and relatives which have seventeen although

this is in part because of the currently unresolved nature of relationships between the

main clades of core eudicots (this is likely to change soon) The Main Tree could have

been drawn with Amborella Acorus or a host of other taxa at the far right without

offending any relationships phylogenetic trees are like mobiles the only fixed points

being the nodes Despite the problems just mentioned pectination interpreted

carefully has its value and it is not an arbitrary process As one reads the terminals of a

pectinate tree from left to right adjacent terminals will be separated by apomorphies

that will cumulate in an appropriate fashion When thinking of a book or a herbarium

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

23

sequence (see Haston et al 2007) this is of some value Nymphaeales and

Austrobaileyales are here adjacent on the tree but they could be separated by

hundreds of families in the sequence merely by rotating only the node from which

Nymphaeales and their sister group (all other angiosperms minus about 75 species)

arise If adjacent in a book or herbarium or as in these summary pages then it is

relatively easy to relate their characters both apomorphic and plesiomorphic but if

separated by hundreds of pages or two floors in a large building then it is less easy to

get anything from the sequence (Since all orders in the pages of the main part of this

site are preceded by the apomorphies of all nodes immediately below them in the seed

plant phylogeny and because of the linkages that have been built in to the site this

problem is much less serious there there is no sequence) So the particular sequence of

all seed plants that is based on a phylogenetic classification but is to be used in herbaria

or any other place where taxa have to be arranged linearly as in this part of APweb

will be one that maximizes the number of taxa that are both successive branches of the

tree and placed successively in the sequence Since specimens are generally filed under

families the outline of a new family sequence for arranging herbaria and books can now

be suggested (Haston et al 2007) and such a standardized sequence will further help

teaching and learning about plants

ON THINKING ABOUT APOMORPHIES

Identifying apomorphies is important because understanding the evolution of

morphology in the broad sense - ie including anatomy chemistry etc - in the context

of diversification is one of our major goals For identifying apomorphies several

preconditions must be met One needs to have an accurate well supported phylogeny

one has to have examined the right taxa both from the point of view of morphology and

molecules one has to have coded the characters right (ie delimited states

appropriately) and one has to use an appropriate model of evolution when fitting the

variation to the tree (see eg Omland 1999 Stevens 2006b) If the distributions of

apomorphies below are compared with those in other studies (eg Turgeon et al 2001

Bremer et al 2001 Endress 2001 Albach et al 2001a Judd amp Olmstead 2004 D Soltis

et al 2005b Zhang et al 2006 Judd et al 2007) the reader may find a number of

differences Although I have integrated such studies as far as is possible there are five

reasons that link to the preconditions just mentioned why there may be differences

where features are placed on trees

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

24

1 I may not have found all the information about a particular character there may be

disagreement over its interpretation or I have added information

2 The sampling of nearly all molecular studies is very incomplete (see Salisbury amp Kim

2001 for problems caused by sampling) indeed it is commonplace to decry the

incompleteness of molecular sampling But not only is the sampling in molecular studies

often less than we might wish that of the morphological and chemical characters whose

evolution we are interested in understanding is also often very poor So for many

anatomical chemical and embryological characters that are confidently said to

characterise families and other groups we all too often have no idea if those characters

are applicable to the whole clade or just to a subgroup within that clade However as

our sampling improves we can locate changes on the tree more precisely if in

unexpected positions Thus is is sometimes suggested that Ericaceae have ellagic acid

(Soltis et al 2005b) in fact the little evidence we have indicates that only a small clade

of some 80 species has ellagic acid and the rest of the family - itself alone about one

third of all Ericales - does not Similarly Ericaceae are noted for their

ectendomycorrhizal associations but Enkianthus sister to all other Ericaceae

apparently lacks these (Abe 2005) Thus these mycorrhizae and their associated

distinctive hair roots are unlikely to be apomorphies of Ericaceae - however it appears

that other families near Ericaceae may also have ectendomycorrhizae (see Asai 1934)

3 I am fitting characters to a very conservative tree with many polytomies although the

nodes that are utilised are for the most part strongly supported polytomies make the

optimisation of characters that is the assigment of character state change to a

particular node on the tree notably difficult (eg Madison amp Madison 2002) In nearly

all studies of the evolution of characters distributions of characters are optimised on a

more or less fully resolved tree and the construction of supertrees may yield yet more

detailed hypotheses of relationships (for literature on supertrees see Cotton amp

Wilkinson 2007 2008) Of course some nodes on such fully resolved trees andor

supertrees may have little support and optimisations of characters on such trees may

carry correspondingly little conviction Even parts of some of the trees used here have

poor support eg relationships within aquatic Alismatales etc although I indicate

when this is the case

4 Exactly how one goes about optimising a character on a tree is critically important

Even using simple parsimony optimisations (ACCTRAN or DELTRAN ACCelerated

TRANsitions or DELayed TRANsitions) the position of synapomorphies on trees - and

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

25

hence our ideas of evolution - may differ and this is still more true if one uses maximum

likelihood or Bayesian methods and makes apparently reasonable suggestions about

weighting gains over losses (eg Donoghue amp Ackerley 1996 Cunningham et al 1998

Omland 1997 1999 Ree amp Donoghue 1999 Polly 2001 Webster amp Purvis 2001

Ronquist 2004 Crisp amp Cook 2005) Sannier et al (2007) show how in in Arecaceae that

where on a phylogenetic tree one might peg changes in microsporogenesis will depend

on the methods one uses to do this and Pedersen et al (2007) discuss the sometimes

very substantial effect of node support on the posterior probabilities of ancestral

character states Here I use parsimony optimization not always as explicit as it might be

but I have often indicated where there are particularly important uncertainties as to the

positions of particular character changes on the tree

5 Finally although I have paid quite a lot of attention to the delimitation of the

character states that make up all the characterizations I have not spent enough time on

this critical operation If we are interested in understanding evolution then fitting the

basic variation - not character states - to a tree in principle allows greater flexibility in

understanding morphology in the context of local phylogenies (see also Stevens 2000

Endress 2005c) However many character states used here are delimited globally that

is they are circumscribed in the context of the variation shown by individual characters

across all angiosperms andor in the context of classic ideas of character evolution

Character states often have arbitrary limits and serve best to communicate

information whether they are in fact suitable for either phylogenetic analysis or

understanding evolution are separate issues Studies have rather unsurprisingly

perhaps but importantly shown that dividing the one character into different sets of

states may yield differing ideas of evolution of that character (eg Lamb Frye amp Kron

2003 Hibbett 2004) When looking at trees on which character states are optimised

one should bear in mind the problems surrounding the delimitation of states (eg

Stevens 2000 2006b) and the danger of using pollen or other types - constructs

based on many characters that vary independently but which effectively get lost in

these types thus Blackmore et al (2009) decomposed the pollen types in Asteraceae

into 52 characters

I use Remanes three main criteria of homology or better similarity when

determining the basic similarity of structures on different organisms (see Remane 1952)

These criteria are special properties position and intermediates Special properties

include anatomical or chemical characters gene expression data etc Position refers

to the position of an organ with respect to landmarks on the plant Although plants are

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

26

plants and landmarks are not as fixed as one might like trying to work out the relative

positions of parts is a good way of understanding morphology so long as one does not

try to out-do Procrustes Intermediates include intermediates found on the same

plant or on different plants Intermediates may observed during development or be

adult structures Thus one may find changes during development which suggest what

the nature of a particular structure is and very different stuctures on different plants

may turn out to be similar early in development Or one can simply compare different

taxa and see that structures that appear to be very different when comparing two

different taxa can be linked morphologically by looking at other taxa

Nevertheless the use of these criteria may not yield an unambiguous answer as to

what a structure is even given a solid phylogeny and an improved understanding of

development (see Jaramillo amp Kramer 2007 for a useful discussion) As Endress (2005c)

observed a number of features - position function development shape anatomy

histology gene activity and relationships to other taxa that clearly have petals - can be

used to distinguish a petal (for example) from other floral structures if a petal does not

have one of these features is it thereby not a petal Thus Maturen et al (2005) recently

found that floral organ diversity genes (B and C) were expressed in the large white

inflorescence bracts of Cornus (see also Costa et al 2005) Peney et al (2005) noted that

not all monosulcate pollen grains in monocots have the same developmental pathway

and that as a result such pollen might not have the same ancestral state Reeves and

Olmstead (2003) suggested that the genetic mechanisms causing monosymmetry in

Lamiales and Solanales were different and Serna and Martin (2006) described similar

problems with the development of hairs in Arabidopsis when compared with that of

hairs in Antirrhinum and Solanaceae Indeed as one perhaps might expect delimitation

of states and characters does not necessarily become easier with increasing knowledge

of development etc Thus Buzgo et al (2004) Matthews and Endress (2005) and others

have shown how hard it can be to distinguish between eg prophylls and other floral

structures as their behaviour is studied during the course of the development of the

flower Similarly at what concentration is a particular secondary metabolite deemed to

be present (Waterman (2007)

The validity of the approach used here that of fitting morphological variation to a

largely molecular-based tree may be questioned However I think it rather unlikely that

well-supported molecular branches will be overturned by morphological data Indeed

analyses of morphological data alone do provide support for many of the clades evident

in molecule-only analyses and analyses with morphological and molecular data

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

27

together may lead to increased support for clades (eg Hufford 1992 Nandi et al 1998

[but adding morphological data reduces support for a number of critical clades too]

Doyle amp Endress 2000 however in none of these papers is the use of morphology

without ambiguity) It is unfortunately clear that the use of morphology alone may not

suggest problems in the phylogenetic placement of taxa that later turn out to have been

wrongly included (eg Zhang et al 1992) Thus I have been wary of putting much weight

on clades that have only morphological support but note that I have treated molecule-

based clades with low bootstrap or jacknife support values (esp below 70) or low

posterior probabilies (below 095) likewise Although I may have been mistaken in

placing so much emphasis on molecular data in terms of providing the basic

phylogenetic framework for angiosperms morphological and molecular data are only

very rarely in irreconcilably strong conflict There are indeed a few places where the

conflict seems extreme These include the relative positions of the Monimiaceae and

Hernandiaceae (Laurales) the position of Hanguanaceae (Commelinales [as here] or

Zingiberales) and of Triplostegia (is it in Dipsacaceae or Valerianaceae - see

Dipsacales) Fossils are unlikely to affect the topologies of the trees presented here but

see below for their importance in understanding morphological evolution in general and

the evolution of angiosperms in particular Nevertheless some largely reject the idea

that trees based on molecular data alone can recover phylogenetic relationships

especially when branching points are old and prefer to used trees based on analysis of

morphological data including those taken from fossils (Hilton amp Bateman 2006 Farjon

2007)

All in all however the extent of the congruence between morphological and

molecular data is impressive and heartening and many clades can be characterised

morphologically It seemed in 1998 that there were no unambiguous morphological

synapomorphies for angiosperm orders (K Bremer 2000) and this is still true if by

unambiguous is meant non-homoplasious However many orders and other clades

have synapomorphies even if these may be indistinguishable at least at the current

level of morphological and developmental knowledge from parallel occurences

elsewhere As our morphological knowledge increases so too does the number of

apomorphies

Endress and Matthews (2006a) emphasize the importance of tendencies and

developmental constraints when thinking about characters of clades within the rosids -

many characters or character combinations occur in a rather sporadic fashion within

the clade and apparently notably less frequently outside it There are a number of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

28

examples of tendencies here and in the individual order pages such as the features

enclosed by parentheses in groups above the level of family eg see (cuticular waxes

as aggregated rodlets) for the commelinids and the discussion of the distribution of

polyandry that is flowers with numerous stamens in the asterid I + II groups etc

Tendencies may involve more than single characters When thinking of the

associations of features like integument thickness and vascularization of the integument

(Endress amp Matthews 2006a) one can imagine fairly simple developmental

preconditions being operative As Endress and Matthews (2006a) note it is difficult to

imagine vascular tissue developing in an integument only two cells thick Changes here

would be loosely correlated if morphologically linked However the strongly correlated

changes noted by Givnish et al (2005) are ecologically linked but are presumably

morphologicallydevelopmentally independent When there seem to be characters

evolving more or less together tests can be carried out to see if the changes are

concentrated on certain branches of the tree (eg Maddison 1990 Sanderson 1991

Maddison amp Maddison 2000) Such tests have rarely been carried out at higher levels in

angiosperms

One of the most striking examples of a tendency is the distribution of N-fixation

restricted as it is to a monophyletic group of four clades although it has arisen seven (or

perhaps several more) times independently within the N-fixing clade and several

members of two quite different kinds of bacteria are involved (eg Clawson et al 2004

Elliott et al 2007 Sprent amp James 2007) Other examples of work that bears on the issue

of tendencies include the findings that flowers of polysymmetrical Arabidopsis have

genes like TCP1 that are expressed asymmetrically during early development and TCP1

is a probable orthologue of the well-known CYC gene of Antirrhinum that is involved in

the development of monosymmetric slowers there (Cubas et al 2001 Costa et al 2005

etc) Parallelism might build on this underlying morphologically cryptic monosymmetry

even if details of the genetic mechanisms causing the monosymmetry evident in

particular groups may be different (Reeves amp Olmstead 2003 Cubas 2004 see above)

Similarly there have been several recent suggestions that the capability to synthesise a

particular metabolite may be switched off but not lost and so can sometimes be

reacquired (eg Wink amp Witte 1983 Wink 2003 Liscombe et al 2005 Larsson 2007

Waterman 2007) Hence perhaps the rather spotty distribution of many secondary

metabolites like ellagic acid the indole alkaloid camptothecin iridoids etc when

considered in the context of phylogenies As a non-botanical example - but a rather nice

one - Salwini-Plawen and Mayr (1961) suggested some time ago that there has been

considerable parallelism (40-65 or more independent origins) in the evolution of eyes in

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

29

metazoans However Pax 6 seems to be a master control gene that is involved in eye

formation perhaps throughout all bilateralians (eg Gehring amp Ikeo 1999 Erwin amp

Davidson 2002) which changes the nature of the problem Note however that in

general our invocation of evolutionary tendencies to explain the patterns of

distributions of characters is really an admission that we do not understand those

patterns

As we find out more about variation we will probably find fewer and fewer features

constant throughout a group Most unqualified statements of presence and absence

should properly be qualified as usually present or usually absent if one is thinking of

the characterisations on the order pages as encompassing the total variation within a

clade Thus Pistia alone among monocots as so far known has sieve tube plastids with

starch grains not protein crystals However this does not affect the fact that sieve tube

plastids with cuneate protein crystals and lacking starch grains are an apomorphy for

monocots Furthermore fossils in a number of cases suggest character combinations

unknown in extant taxa as may be seen in the discussions of Fagaceae Platanaceae

Iteaceae Calycanthaceae etc Confusing the issue there may be questions as to where

exactly on the tree a particular fossil is to be placed (eg see Nymphaeaceae

Calycanthaceae Archaefructus etc)

To summarize given our current understandings of both phylogenies and

characters evolution of some characters in which we are interested seems very labile

(see eg D Soltis et al 2005b Endress and Matthews 2006a Stevens 2006b Ekman et

al 2008 for a good recent study) and I have been cautious when talking about character

evolution Much effort must continue to be spent in summarizing characters of well-

established clades at all levels providing features by which they may be recognized and

signaling synapomorphies Remember that (1) the basic morphological anatomical and

chemical knowledge of many critical taxa is woefully incomplete (2) different

assumptions about character evolution may greatly affect the position of

synapomorphies on trees (3) in many cases relationships within and between many

groups are too uncertain at present to worry very much about synapomorphies and

(4) we must be clear about what we do and do not not know As mentioned above I

have indicated in a number of places where there are particularly important

uncertainties as to where characters should be placed on the tree Nevertheless it is a

relatively easy matter to update notes such as these and it can be a simple matter to

incorporate new data on characters that have never before been considered in the

context of a tree Much basic - and unfortunately perhaps unfashionable - work must be

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

30

carried to clarify the distribution of morphological anatomical and chemical characters

for examples of what can to be done see the work of P K Endress and collaborators

(floral morphology and development) and S R Jensen and collaborators (iridoids) But

acquiring information about nodal anatomy cell and tissue distribution nodal anatomy

and the like is not difficult at all and should be generally encouraged

But all this is merely a necesary prelude to the understanding of evolution For this

one needs to know a lot more including dating the phylogeny understanding the

function(s) of characters factoring in relevant aspects of the palaeoenvironment in

which an apomorphy first appeared etc

SUMMARY OF THE SYSTEM

Below is a formalised summary of the relationships within orders of the families of seed

plants There are a few families that are not recognised even as options in APG II and

vice versa as well as a few extra orders The families to be recognised in the most

recent edition of Mabberleys The Plant Book (Mabberley 2008) are also largely

consistent with those below But all differences are trivial and will - I hope - eventually

disappear

Square brackets - [] - enclose clades the plus sign - + - designates sister taxa a

comma - - denotes part of a polytomy and quotation marks - - denotes a

paraphyletic group

SEED PLANTS

GYMNOSPERMS

Cycadales

Cycadaceae + Zamiaceae

Ginkgoales

Ginkgoaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

31

Pinales

Pinaceae [[Araucariaceae [Podocarpaceae + Phyllocladaceae]] [Sciadopityaceae

[Taxaceae + Cupressaceae]]]

Gnetales

Ephedraceae [Gnetaceae + Welwitschiaceae]

ANGIOSPERMSFLOWERING PLANTS

Amborellales

Amborellaceae

Nymphaeales

Hydatellaceae [Cabombaceae + Nymphaeaceae]

Austrobaileyales

Austrobaileyaceae [Schisandraceae + Trimeniaceae]

Chloranthales

Chloranthaceae

MAGNOLIIDS

Magnoliales

Myristicaceae [Magnoliaceae [[Himantandraceae + Degeneriaceae] [Eupomatiaceae +

Annonaceae]]]

Laurales

Calycanthaceae [[Siparunaceae [Gomortegaceae + Atherospermataceae]] [Monimiaceae

[Hernandiaceae + Lauraceae]]]

Canellales

Canellaceae + Winteraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

32

Piperales

[Hydnoraceae + Aristolochiaceae] [Piperaceae + Saururaceae]

MONOCOTS

Acorales

Acoraceae

Alismatales

Araceae [Tofieldiaceae [[Alismataceae [Hydrocharitaceae + Butomaceae]]

[Scheuchzeriaceae [Aponogetonaceae [Juncaginaceae [Maundiaceae [[Posidoniaceae

[Ruppiaceae + Cymodoceaceae]] [Zosteraceae + Potamogetonaceae]]]]]]]

Petrosaviales

Petrosaviaceae

Dioscoreales

Nartheciaceae [[Taccaceae + Thismiaceae] [Burmanniaceae + Dioscoreaceae]]

Pandanales

Velloziaceae Triuridaceae Stemonaceae [Pandanaceae + Cyclanthaceae]

Liliales

Corsiaceae [Campynemataceae [Petermanniaceae [Colchicaceae + Alstroemeriaceae]]

Melanthiaceae [[Philesiaceae + Rhipogonaceae] [Smilacaceae + Liliaceae]]]

Asparagales

Orchidaceae [[Boryaceae [Blandfordiaceae [Lanariaceae [Asteliaceae + Hypoxidaceae]]]]

[[Ixioliriaceae + Tecophilaeaceae] [Doryanthaceae [Iridaceae [Xeronemataceae

[Xanthorrhoeaceae [Alliaceae + Asparagaceae]]]]]]]

COMMELINIDS

Unplaced

Dasypogonaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

33

Arecales

Arecaceae

Poales

[Typhaceae + Bromeliaceae] [Rapateaceae [[Thurniaceae [Juncaceae + Cyperaceae]]

[[Mayacaceae [Eriocaulaceae + Xyridaceae]] [Flagellariaceae [Anarthriaceae

[Centrolepidaceae + Restionaceae]]] [Joinvilleaceae [Ecdeiocoleaceae + Poaceae]]]]]]]

Commelinales

[Commelinaceae + Hanguanaceae] [Philydraceae [Haemodoraceae + Pontederiaceae]]

Zingiberales

Musaceae [Strelitziaceae + Lowiaceae] Heliconiaceae [[Cannaceae + Marantaceae]

[Costaceae + Zingiberaceae]]

Ceratophyllales

Ceratophyllaceae

EUDICOTS

Ranunculales

Eupteleaceae [Papaveraceae [[[Lardizabalaceae + Circaeasteraceae] [Menispermaceae

[Berberidaceae + Ranunculaceae]]]]

Sabiales

Sabiaceae

Proteales

Nelumbonaceae [Platanaceae + Proteaceae]

Trochodendrales

Trochodendraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

34

Buxales

Haptanthaceae [Buxaceae + Didymelaceae]

CORE EUDICOTS

Gunnerales

Gunneraceae + Myrothamnaceae

Dilleniales

Dilleniaceae

Saxifragales

Peridiscaceae [[Paeoniaceae [Altingiaceae [Hamamelidaceae [Cercidiphyllaceae +

Daphniphyllaceae]]]] [[Crassulaceae [Aphanopetalaceae [Tetracarpaeaceae

[Penthoraceae + Haloragaceae]]]] [Iteaceae [Grossulariaceae + Saxifragaceae]]]]

Cynomoriaceae unplaced

Vitales

Vitaceae

ROSIDS

FABIDROSID I

Zygophyllales

Krameriaceae + Zygophyllaceae

Celastrales

Lepidobotryaceae + Celastraceae

Oxalidales

Huaceae [[Connaraceae + Oxalidaceae] [Cunoniaceae [Elaeocarpaceae [Brunelliaceae +

Cephalotaceae]]]]

Malpighiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

35

[Achariaceae [Goupiaceae [Violaceae + Passifloraceae] [Lacistemataceae Salicaceae]]

[Lophopyxidaceae Putranjivaceae] [Ctenolophonaceae [Erythroxylaceae +

Rhizophoraceae]] Linaceae Ixonanthaceae Humiriaceae Irvingiaceae

Centroplacaceae Pandaceae [[Bonnetiaceae + Clusiaceae] [Calophyllaceae

[Hypericaceae + Podostemaceae]]] [Malpighiaceae + Elatinaceae] Ochnaceae

[Peraceae [Rafflesiaceae + Euphorbiaceae]] [Phyllanthaceae + Picrodendraceae]

[Balanopaceae [[Trigoniaceae + Dichapetalaceae] [Chrysobalanaceae +

Euphroniaceae]]] Caryocaraceae

N-FIXING CLADE

Fabales

Quillajaceae [Fabaceae [Polygalaceae + Surianaceae]]

Rosales

Rosaceae [[Barbeyaceae [Elaeagnaceae Dirachmaceae Rhamnaceae]] [Ulmaceae

[Cannabaceae [Moraceae + Urticaceae]]]]

Cucurbitales

Anisophylleaceae [[Corynocarpaceae + Coriariaceae] [Cucurbitaceae [Tetramelaceae

[Datiscaceae + Begoniaceae]]]] Apodanthaceae

Fagales

Nothofagaceae [Fagaceae [[Myricaceae + Juglandaceae] [Casuarinaceae

[Ticodendraceae + Betulaceae]]]]

MALVIDROSID II

pgtGeraniales

Geraniaceae [[Melianthaceae + Francoaceae] [Vivianaceae + Ledocarpaceae]]

Myrtales

Combretaceae [[Onagraceae + Lythraceae] [[Vochysiaceae + Myrtaceae]

[Melastomataceae [Crypteroniaceae [Alzateaceae + Penaeaceae]]]]]

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

36

Crossosomatales

[Staphyleaceae [Guamatelaceae [Crossosomataceae + Stachyuraceae]]] [Aphloiaceae

[Geissolomataceae + Strasburgeriaceae]]

Picramniales

Picramniaceae

Sapindales

Biebersteiniaceae Nitrariaceae [Kirkiaceae [Anacardiaceae + Burseraceae]]

[Sapindaceae [Simaroubaceae Rutaceae Meliaceae]]

Huerteales

Gerradinaceae [Dipentodontaceae + Tapisciaceae]

Malvales

Neuradaceae [Thymelaeaceae [Sphaerosepalaceae Bixaceae [Cistaceae

[Sarcolaenaceae + Dipterocarpaceae]] [Cytinaceae + Muntingiaceae] Malvaceae]]

Brassicales

[Akaniaceae + Tropaeolaceae] [[Moringaceae + Caricaceae] [Setchellanthaceae

[Limnanthaceae [[Koeberliniaceae [Bataceae + Salvadoraceae]] [Emblingiaceae

[Pentadiplandraceae [Gyrostemonaceae Resedaceae Stixis and relatives] Tovariaceae

[Capparaceae [Cleomaceae + Brassicaceae]]]]]]]]

Berberidopsidales

Aextoxicaceae + Berberidopsidaceae

Santalales

Erythropalaceae Olacaceae [[Loranthaceae [Misodendraceae + Schoepfiaceae]]

[Opiliaceae + Santalaceae]] Balanophoraceae unplaced

Caryophyllales

[[Droseraceae [Nepenthaceae [Drosophyllaceae [Ancistrocladaceae +

Dioncophylleaceae]]]]] [[Frankeniaceae + Tamaricaceae] [Polygonaceae +

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

37

Plumbaginaceae]]] [Rhabdodendraceae [Simmondsiaceae [[Asteropeiaceae +

Physenaceae] [[Caryophyllaceae [Achatocarpaceae + Amaranthaceae]]

[Stegnospermataceae [Limeaceae [[Lophocarpaceae [Barbeuiaceae [Aizoaceae

[Sarcobataceae Phytolaccaceae Nyctaginaceae]]]] [Molluginaceae [Halophytaceae

Montiaceae Basellaceae Didiereaceae [Talinaceae [Portulacaceae [Anacampseros etc +

Cactaceae]]]]]]]]]]]]]

ASTERIDS

Cornales

[Cornaceae + Nyssaceae] [Hydrangeaceae + Loasaceae] Hydrostachyaceae [Curtisiaceae

+ Grubbiaceae]

Ericales

[Balsaminaceae [Marcgraviaceae + Tetrameristaceae]] [[Polemoniaceae +

Fouquieraceae] Lecythidaceae [[Sladeniaceae + Pentaphylacaceae] [Sapotaceae

[Ebenaceae [Maesaceae [Theophrastaceae [Myrsinaceae + Primulaceae]]]]]

[Mitrastemonaceae Theaceae [Symplocaceae [Styracaceae + Diapensiaceae]]

[[Sarraceniaceae [Roridulaceae + Actinidiaceae] [Clethraceae [Cyrillaceae +

Ericaceae]]]]]]

LAMIIDASTERID I

Unplaced

Metteniusaceae Oncothecaceae Icacinaceae and some unplaced ex-Icacinaceae

Garryales

Garryaceae + Eucommiaceae

Unplaced

Boraginaceae Vahliaceae

Gentianales

Rubiaceae [Gentianaceae [Loganiaceae [Gelsemiaceae + Apocynaceae]]]

Lamiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

38

Plocospermataceae [Oleaceae [Tetrachondraceae [Calceolariaceae

[Gesneriaceae Peltanthera [Pedaliaceae Carlemanniaceae Martyniaceae Stilbaceae

Plantaginaceae Linderniaceae Bignoniaceae Verbenaceae Lamiaceae Paulowniaceae

Schlegeliaceae Thomandersiaceae Phrymaceae Rehmannia Acanthaceae

Scrophulariaceae Orobanchaceae Byblidaceae Lentibulariaceae]]]]]

Solanales

[Montiniaceae [Sphenocleaceae + Hydroleaceae]] [Convolvulaceae + Solanaceae]

CAMPANULIDASTERID II

Aquifoliales

[[Cardiopteridaceae + Stemonuraceae] [Aquifoliaceae [Helwingiaceae +

Phyllonomaceae]]

Asterales

[Rousseaceae + Campanulaceae] Pentaphragmataceae [[Alseuosmiaceae [Phellinaceae

+ Argophyllaceae]] Stylidiaceae [Menyanthaceae [Goodeniaceae [Calyceraceae +

Asteraceae]]]]

Escalloniales

Escalloniaceae

Bruniales

[Bruniaceae + Columelliaceae]

Apiales

Pennantiaceae [Torricelliaceae [Griseliniaceae [Pittosporaceae [Araliaceae

[Myodocarpaceae + Apiaceae]]]]]

Paracryphiales

Paracryphiaceae

Dipsacales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

39

Adoxaceae [Diervillaceae [Caprifoliaceae [Linnaeaceae [Morinaceae [Dipsacaceae +

Valerianaceae]]]]]

LINKS TO ORDERS AND FAMILIES

[Back to Top]

MAIN GROUPINGS

Asterids commelinids core eudicots asterid 1 asterid

2 eudicots gymnosperms Magnoliophyta monocots N-fixing claderosids seed plants

ALPHABETICAL LISTING OF ALL ORDINAL NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | L | M | N | O | P | Q | R | S | T | U | V | W | X | Z

Abietales Acanthales Acerales Acorales Actinidiales Actinostrobales Adoxales Aescul

ales Agavales Aizoales AkanialesAlismatales Alliales Alseuosmiales Alstroemeriales

Altingiales Amaranthales Amaryllidales Amborellales AmbrosialesAmmiales Amomal

es Ancistrocladales Anisophylleales Annonales Anthobolales Apiales Apocynales Apo

nogetonalesAquifoliales Arales Araliales Aralidiales Araucariales Arecales Aristolochi

ales Athrotaxidales Asarales AsclepiadalesAsparagales Asphodelales Asparagales Ast

eliales Atriplicales Aucubales Austrobaileyales Avenales

Balanitales Balanopales Balanophorales Balsaminales Barbeyales Barclayales Batales

Begoniales BerberidalesBerberidopsidales Betulales Biebersteiniales Bignoniales Bix

ales Boraginales Brassicales Brexiales Bromeliales BrunialesBrunoniales Burmannial

es Burserales Butomales Buxales Byblidales

Cactales Callitrichales Calycanthales Calycerales Campanulales Campynematales Can

ellales Cannales CapparalesCaprifoliales Cardiopteridales Carduales Caricales Carle

manniales Caryophyllales Cassiales Casuarinales CelastralesCentrolepidales Cephalot

ales Cephalotaxales Ceratophyllales Cercidiphyllales Chenopodiales Chironiales Chlor

anthalesChrysobalanales Cinchonales Circaeasterales Cistales Citrales Cocosales Col

chicales Columelliales CombretalesCommelinales Connarales Convolvulales Coriarial

es Cornales Corylales Corynocarpales Crassulales CrossosomatalesCucurbitales Cunn

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

40

inghamiales Cunoniales Cupressales Cycadales Cyclanthales Cymodoceales Cynarales

CynomorialesCyperales Cyrillales Cytinales

Daphnales Daphniphyllales Dasypogonales Datiscales Degeneriales Desfontainiales D

iapensiales Didymelales DillenialesDioales Dioncophyllales Dioscoreales Diospyrales

Dipentodontales Dipsacales Droserales

Ebenales Echiales Elaeagnales Elaeocarpales Elatinales Elodeales Emmotales Empetr

ales Ephedrales Ericales EriocaulalesErythropalaless Escalloniales Eucommiales Euph

orbiales Eupomatiales Eupteleales Euryalaless

Fabales Fagales Falcatifoliales Ficales Flacourtiales Flagellariales Fouquieriales Franc

oales Frangulales

Galiales Garryales Geissolomatales Gentianales Geraniales Gesneriales Ginkgoales G

laucidiales Globulariales GnetalesGoodeniales Greyiales Griseliniales Grossulariales

Grubbiales Gunnerales Gyrocarpales Gyrostemonales

Haemodorales Haloragales Hamamelidales Hanguanales Heisteriales Helleborales He

lwingiales Himantandrales HippuridalesHomaliales Hortensiales Huales Huerteales

Hydatellales Hydnorales Hydrangeales Hydrastidales HydrocharitalesHydropeltidales

Hydrostachyales Hypericales Hypoxidales

Icacinales Illiciales Iridales Irvingales Iteales Ixiales

Jasminales Juglandales Julianiales Juncaginales Juncales

Lacistematales Lactoridales Lamiales Lardizabalales Laurales Lecythidales Ledocarpal

es Leitneriales LentibularialesLigustrales Liliales Limnanthales Linales Liriales Loasal

es Lobeliales Loganiales Lonicerales Loranthales LowialesLythrales

Magnoliales Malpighiales Malvales Marathrales Marcgraviales Mayacales Medusagy

nales Medusandrales MelanthialesMelastomatales Meliales Melianthales Meliosmal

es Menispermales Menyanthales Metteniusales MitrastemonalesMiyoshiales Moni

miales Moringales Musales Myricales Myristicales Myrothamnales Myrsinales Myrta

les

Najadales Nandinales Narcissales Nartheciales Nelumbonales Nepenthales Neuradal

es Nitrariales Nolanales NothofagalesNyctaginales Nymphaeales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

41

Ochnales Oenotherales Olacales Oleales Onagrales Oncothecales Opuntiales Orchid

ales Oxalidales

Saxifragales Pandales Pandanales Papaverales Paracryphiales Parasitaxales Paridales

Parnassiales PassifloralesPenaeales Pennantiales Pentaphragmatales Peridiscales P

etiveriales Petrosaviales Phellinales Philydrales PhyllanthalesPhysenales Phytolaccal

es Picramniales Pinales Pinguiculales Piperales Pittosporales Plantaginales Platanale

sPlumbaginales Poales Podocarpales Podophyllales Podostemales Polemoniales Pol

ygalales Polygonales PontederialesPortulacales Posidoniales Potamogetonales Primu

lales Proteales

Quercales Quillajales Quintiniales

Rafflesiales Ranunculales Rapateales Resedales Restionales Rhabdodendrales Rhamn

ales Rhinanthales RhizophoralesRhodorales Rhoipteleales Roridulales Rosales Rouss

eales Rubiales Ruppiales Rutales

Sabiales Salicales Salvadorales Samolales Samydales Sanguisorbales Santalales Sapin

dales Sapotales SarracenialesSaxegotheales Saxifragales Scheuchzeriales Sciadopitya

les Scleranthales Scrophulariales Scyphostegiales SedalesSilenales Simmondsiales S

milacales Solanales Sphenocleales Sphenostemonales Stangeriales Stellariales Stemo

nalesStilbales Stylidiales Styracales Surianales

Taccales Tamales Tamaricales Taxales Taxodiales Tecophilaeales Terebinthales Tern

stroemiales Theales TheligonalesThymelaeales Tiliales Tofieldiales Torricelliales Tov

ariales Tribelales Trilliales Trimeniales Triuridales TrochodendralesTropaeolales Turn

erales Typhales

Ulmales Urticales

Vacciniales Vahliales Vallisneriales Velloziales Veratrales Verbenales Viburnales Vinc

ales Violales Viscales VitalesVochysiales

Welwitschiales Winterales

Xanthorrhoeales Ximeniales Xyridales

Zamiales Zingiberales Zosterales Zygophyllales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

42

ALPHABETICAL LISTING OF ALL FAMILY NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | K | L | M | N | O | P | Q | R | S | T | U | V | W | X | Y

|Z

Back to Home

In some cases clicking on a family synonym will send you directly to the subfamily in

which the genus of the basionym is to be found

Abietaceae Abolbodaceae Abrophyllaceae Acalyphaceae Acanthaceae [Acanthochla

mydaceae] Acarnaceaee AceraceaeAchariaceae Achatocarpaceae Achradaceae Achy

ranthaceae Acmopylaceae Acoraceae Acristaceae ActaeaceaeActinidiaceae Actinota

ceae Amentotaxaceae Adenogrammaceae Adoxaceae Aegialitidaceae Aegicerataceae

Aegilopaceae Aeginetiaceaee Aegiphilaceae Aesculaceae Aextoxicaceae Agapanthac

eae Agavaceae AgdestidaceaeAgialidaceae Agrimoniaceae Agrostidaceae Ailanthace

ae Aitoniaceae Aizoaceae Akaniaceae Alangiaceae AlchemillaceaeAldrovandaceae A

lismataceae Alliaceae Allioniaceae Allophylaceae Aloaceae Alopecuraceae Alpiniacea

e AlseuosmiaceaeAlsinaceae Alsodeiaceae Alstroemeriaceae Altingiaceae Alzateacea

e Amaranthaceae Amaryllidaceae Amborellaceae Ambrosiaceae Amentotaxaceae A

mmanniaceae Ammiaceae Amomaceae Ampelidopsaceae Amygdalaceae Amyridacea

eAnacardiaceae Anagallidaceae Anarthriaceae Anchusaceae Ancistrocladaceae Andr

omedaceae AndropogonaceaeAndrostachyaceae Androsynaceae Anemarrhenaceae

Anemonaceae Angelicaceae Anisophylleaceae AnnonaceaeAnomochloaceae Anopter

aceae Anrederaceae Anthemidaceae Anthericaceae Anthobolaceae Antidesmataceae

Antirrhinaceae Antoniaceae Aparinaceae Aphanopetalaceae Aphloiaceae Aphyllanth

aceae Apiaceae ApocynaceaeApodanthaceae Aponogetonaceae Aporusaceae Apose

ridaceae Apostasiaceae Aptandraceae AquifoliaceaeAquilariaceae Aquilegiaceae A

raceae Aragoaceae Araliaceae Aralidiaceaee Araucariaceae Arbutaceae Arceuthidace

aeArceuthobiaceae Arctostaphylaceae Arctotidaceae Ardisiaceae Arecaceae Argoph

yllaceae Arisaraceae AristolochiaceaeAristoteliaceae Arjonaceae Armeriaceae Artem

isiaceae Arthrotaxidaceae Artocarpaceae Arundinaceae ArundinellaceaeAsaraceae A

sclepiadaceae Ascyraceaee Asiraceae Asparagaceae Asperulaceae Asphodelaceae As

pidistraceaeeAsteliaceae Asteraceae Asteranthaceae Asteropeiaceae Astragalaceae

Astrocarpaceae AthanasiaceaeAtherospermataceae Atriplicaceae Atropaceae Aucuba

ceae Aurantiaceae Austrobaileyaceae Austrotaxaceae AvenaceaeAverrhoaceae Avet

raceae Avicenniaceae Azaleaceae Azimaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

43

Balanitaceae Balanopaceae Balanophoraceae Balsameaceae Balsaminaceae Bambusa

ceae Barbaceniaceae BarbeuiaceaeBarbeyaceae Barclayaceae Barringtoniaceae Base

llaceae Bataceae Baueraceae Bauhiniaceae Baxteriaceae BegoniaceaeBehniaceae B

elangeraceae Belloniaceae Bembiciaceae Berberidaceae Berberidopsidaceae Berryac

eaee BersamaceaeBertyaceae Berzeliaceae Besleriaceae Betaceae Betulaceae Biebe

rsteiniaceae Bifariaceae Bignoniaceae BischofiaceaeBixaceae Blakeaceae Blandfordia

ceae Blattiaceae Blepharocaryaceae Blitaceae Blyxaceae Boerlagellaceae Bolivaracea

eBombacaceae Bonnetiaceae Bontiaceae Boopidaceae Boraginaceae Borassaceae B

oroniaceae BoryaceaeBotryodendraceae Bougainvilleaceae Boweniaceae Brassicacea

e Brachycaulaceae Bretschneideraceae BrexiaceaeBromeliaceae Brownlowiaceae Br

unelliaceae Bruniaceae Brunoniaceae Brunsvigiaceae Bryoniaceae BuchneraceaeBuci

daceae Buddlejaceae Buglossaceae Bulbocodiaceae Bumeliaceae Bupleuraceae Burc

hardiaceae BurmanniaceaeBurseraceae Butneriaceae Butomaceae Buxaceae Byblida

ceae Byttneriaceae

Cabombaceae Cacaoaceae Cactaceae Caesalpiniaceae Caladiaceae Calamaceae Calce

olariaceae CalectasiaceaeCalendulaceae Callaceae Callicomaceae Calligonaceae Calli

traceae Callitrichaceae Calochortaceae CalophyllaceaeCalthaceae Calycanthaceae Ca

lyceraceae Cambogiaceae Camelliaceae Campanulaceae CampynemataceaeCanacom

yricaceae Canellaceae Cannabaceae Cannaceae Canopodaceae Canotiaceae Cansjera

ceae CapparaceaeCaprariaceae Caprifoliaceae Cardiopteridaceae Carduaceae Carica

ceae Carissaceae Carlemanniaceae CarpinaceaeCarpodetaceae Cartonemataceae Ca

ryocaraceae Caryophyllaceae Casearieaceae Cassiaceae CassipoureaceaeCassythacea

e Castaneaceae Castelaceae Casuarinaceae Catesbaeaceae Cathedraceae Cecropiace

ae CedraceaeCedrelaceae Celastraceae Celosiaceae Celtidaceae Centaureaceae Cen

trolepidaceae Centroplacaceae CepaceaeCephalanthaceae Cephalotaceae Cephalota

xaceae Cerastiaceae Ceratoniaceae Ceratophyllaceae CerberaceaeCercidiphyllaceae

Cercocarpaceae Cercodiaceae Cereaceae Cerinthaceae Ceroxylaceae Cestraceae Cev

alliaceaeChailletiaceae Chamaedoraceae Chamaemoraceae Chamelauciaceae Chaun

ochitonaceae Chelidoniaceae ChelonaceaeChenopodiaceae Chimonanthaceae Chingi

thamnaceae Chionographidaceae Chiranthodendraceae ChironiaceaeChlaenaceae Ch

loanthaceae Chloranthaceae Chloridaceae Chlorogalaceae Chrysobalanaceae Cicerac

eae CichoriaceaeCimicifugaceae Cinchoniaceae Circaeaceae Circaeasteraceae Cissac

eae Cistaceae Citraceae Clematidaceae CleomaceaeClethraceae Clusiaceae Cneorac

eae Cnestidaceae Cnicaceae Cobaeaceae Cochlospermaceae Cocosaceae Coffeaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

44

Colchicaceae Coleogynaceae Columelliaceae Combretaceae Commelinaceae Comocl

adiaceae Compositae CompsoaceaeConantheraceae Connaraceae Conostylidaceae C

onvallariaceae Convolvulaceae Cordiaceae CoreopsidaceaeCoriandraceae Coriariacea

e Coridaceae Corispermaceae Cornaceae Corokiaceae Coronillaceae Correaceae Cor

rigiolaceaeCorsiaceae Corydalaceae Corylaceae Corynocarpaceae Coryphaceae Cost

aceae Cotyledonaceae CoulaceaeCoutariaceae Coutoubiaceae Crassulaceae Crescen

tiaceae Cressaceae Crinaceae Crocaceae CroomiaceaeCrossosomataceae Crotonacea

e Cruciferae Cryptaceae Crypteroniaceae Cryptocorynaceae CryptomeriaceaeCtenol

ophonaceae Cucurbitaceae Cunninghamiaceae Cunoniaceae Cupressaceae Curcumac

eae Curtisiaceae CuscutaceaeCyananthaceae Cyanastraceae Cynanchaceae Cyanella

ceae Cycadaceae Cyclanthaceae CyclantheraceaeCyclocheilaceae Cydoniaceae Cymo

doceaceae Cynaraceae Cynocrambaceae Cynomoriaceae Cyperaceae CyphiaceaeCyp

hocarpaceae Cypripediaceae Cyrillaceae Cyrtandraceae Cyrtanthaceae Cytinaceae

Dacrycarpaceae Dactylanthaceae Damasoniaceae Daphnaceae Daphniphyllaceae Das

ypogonaceae DatiscaceaeDaturaceae Daucaceae Davidiaceae Davidsoniaceae Decais

neaceae Deeringiaceae Degeneriaceae DelphiniaceaeDendrophthoaceae Columelliac

eae Detariaceae Dialypetalanthaceae Dianellaceae Dianthaceae DiapensiaceaeDicha

petalaceae Dichondraceae Diclidantheraceae Dicrastylidiaceae Dictamnaceae Didiere

aceae DidymelaceaeDidymocarpaceae Diegodendraceae Diervillaceae Digitalidaceae

e Dilatridaceae Dilleniaceae Dioaceae DionaeaceaeDioncophyllaceae Dioscoreaceae

Diosmaceae Diospyraceae Dipentodontaceae Diphylleiaceae Diplolaenaceae Dipsaca

ceaeDipterocarpaceae Dirachmaceae Disanthaceae Diselmaceae Dodonaeaceae Do

mbeyaceae Donatiaceae DorsteniaceaeDortmannaceae Doryanthaceae Drabaceae D

racaenaceae Dracontiaceae Drimyidaceae Droseraceae DrosophyllaceaeDryadaceae

Duabangaceae Duckeodendraceae Dulongiaceae Durantaceae Durionaceae Dysphani

aceae

Ebenaceae Eccremidaceae Ecdeiocoleaceae Echiaceae Echinopaceae Ehretiaceae Ela

eagnaceae ElaeocarpaceaeElegiaceae Elatinaceae Ellisiaceae Ellisiophyllaceae Elodea

ceae Elytranthaceae Embeliaceae Emblingiaceae EmmotaceaeEmpetraceae Encepha

lartaceae Engelhardtiaceae Enhalaceae Epacridaceae Ephedraceae Ephemeranthacea

eEpilobiaceae Epimediaceae Eragrostidaceae Eremolepidaceae Eremosynaceae Erica

ceae Erinaceae EriocaulaceaeEriogonaceae Eriospermaceae Erodiaceae Erycibaceae

Eryngiaceae Erysimaceae Erythroniaceae ErythropalaceaeErythrospermaceae Erythro

xylaceae Escalloniaceae Eschscholziaceae Eucomidaceae Eucommiaceae Eucryphiace

aeEuonymaceae Eupatoriaceae Euphorbiaceae Euphrasiaceae Euphroniaceae Eupo

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

45

matiaceae Eupteleaceae EuryalaceaeEustrephaceae Euthemidaceae Eutocaceae Exb

ucklandiaceae Exocarpaceae

Fabaceae Fagaceae Falcatifoliaceae Ferulaceae Festucaceae Ficaceae Ficoidaceae Fit

zroyaceae FlacourtiaceaeFlagellariaceae Flindersiaceae Foetidiaceae Forestieraceae

Fothergillaceae Fouquieriaceae Fragariaceae FrancoaceaeFrangulaceae Frankeniacea

e Fraxinaceae Fraxinellaceae Freycinetiaceae Fritillariaceae Fuchsiaceae Fumariacea

eFunkiaceae

Gaiadendraceae Galacaceae Galanthaceae Galaxiaceae Galedupaceae Galeniaceae G

aliaceae Garciniaceae GardeniaceaeGardneriaceae Garryaceae Geissolomataceae Ge

itonoplesiaceae Gelsemiaceae Geniostomaceae GentianaceaeGeonomataceae Geosir

idaceae Geraniaceae Gerrardinaceae Gesneriaceae Gethyllidaceae Gilliesiaceae Gina

lloaceaeGinkgoaceae Gisekiaceae Gladiolaceae Glaucidiaceae Glechomaceae Glinace

ae Globulariaceae Gnaphaliaceae GnetaceaeGoetzeaceae Gomortegaceae Gomphiac

eae Gomphrenaceae Gonystylaceae Goodeniaceae Gordoniaceae GouaniaceaeGoup

iaceae Gramineae Gratiolaceae Grewiaceae Greyiaceae Grielaceae Griseliniaceae Gr

onoviaceae GrossulariaceaeGrubbiaceae Guaiacanaceae Guamatelaceae Guettardac

eae Gunneraceae Gustaviaceae Guttiferae GyrocarpaceaeGyrostemonaceae

Hachetteaceae Haemanthaceae Haemodoraceae Halesiaceae Hallieraceae Halocarpa

ceae Halophilaceae HalophytaceaeHaloragaceae Hamamelidaceae Hanguanaceae H

aptanthaceae Harmandaceae Hebenstretiaceae HectorellaceaeHederaceae Hedyosm

aceae Hedyotidaceae Hedysaraceae Heisteriaceae Heleniaceae Heliamphoraceae H

elianthaceaeHelianthemaceae Helichrysaceae Heliconiaceae Helicteraceae Heliotropi

aceae Helleboraceae Heloniadaceae HelosidaceaeHelwingiaceae Hemerocallidaceae

Hemimeridaceae Hemiphylacaceae Henriqueziaceae Henslowiaceae Hermanniaceae

Hernandiaceae Herniariaceae Herreriaceae Hesperocallidaceae Heterantheraceae He

teropyxidaceae HeterostylaceaeHibbertiaceae Hibiscaceae Hilleriaceae Himantandra

ceae Hippocastanaceae Hippocrateaceae HippomanaceaeHippophaeaceae Hippurida

ceae Hirtellaceae Holacanthaceae Homaliaceae Hopkinsiaceae Hoplestigmataceae H

ordeaceaeHornschuchiaceae Hortensiaceae Hortoniaceae Hostaceae Houstoniaceae

Huaceae Huerteaceae HugoniaceaeHumbertiaceae Humiriaceae Hyacinthaceae Hyd

atellaceae Hydnoraceae Hydrangeaceae Hydrastidaceae HydrillaceaeHydrocharitacea

e Hydrocotylaceae Hydrogetonaceae Hydroleaceae Hydropeltidaceae Hydrophylacac

eae HydrophyllaceaeHydrostachyaceae Hymenocardiaceae Hypecoaceae Hyperanth

eraceae Hypericaceae Hypopityaceae HypoxidaceaeHypseocharitaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

46

Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta

ceae ImperatoriaceaeInocarpaceae Inulaceae Ionidiaceae Iriarteaceae Iridaceae Irvi

ngiaceae Isnardiaceae Isophysidaceae IteaceaeStrasburgeriaceae Ixiaceae Ixioliriace

ae Ixonanthaceae

Jalapaceae Jamboliferaceae Japonoliriaceae Jasionaceae Jasminiaceae Johnsoniaceae

Joinvilleaceae JuglandaceaeJulianiaceae Juncaceae Juncaginaceae Juniperaceae Jus

siaeaceae Justiciaceae

Kadsuraceae Kaliphoraceae Kalmiaceae Kaniaceae Kiggelariaceae Kingdoniaceae Kin

giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae

Krameriaceae

Labiatae Lacandoniaceae Lachenaliaceae Lacistemataceae Lactoridaceae Lactucaceae

Lagerstroemiaceaee LamiaceaeLampsanaceae Lanariaceae Langsdorffiaceae Lantan

aceae Lapageriaceae Lardizabalaceae Lasiopetalaceae LauraceaeLawsoniaceae Laxm

anniaceae Lecythidaceae Ledaceae Ledocarpaceae Leeaceae Leguminosae Leitneriac

eae LemnaceaeLennoaceae Lentibulariaceae Lentiscaceae Leoniaceae Leonticaceae

Lepidobotryaceae LepidocarpaceaeLepidocaryaceae Lepidocerataceae Leptaulaceae

Leptospermaceae Lepturaceae Lepuropetalaceae LeuctenbergiaceaeLibocedraceae

Licaniaceae Ligustraceae Lilaeaceae Liliaceae Limeaceae Limnanthaceae Limnocharit

aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace

ae Linderniaceae Lindleyaceae Linnaeaceae LippayaceaeLiriaceae Liriodendraceae Li

ssocarpaceae Littorellaceae Loasaceae Lobeliaceaee Loganiaceae LomandraceaeLoni

ceraceae Lopeziaceae Lophiocarpaceae Lophiolaceae Lophiraceae Lophophytaceae L

ophopyxidaceae LoranthaceaeLotaceae Lowiaceae Lupulaceaee Luxemburgiaceae Lu

zuriagaceae Lychnidaceae Lyciaceae LyginiaceaeLygodisodeaceae Lysimachiaceae Ly

thraceae

Macarisiaceae Mackinlayaceae Madiaceae Maesaceae Magnoliaceae Malaceae Male

sherbiaceae MalortieaceaeMalpighiaceae Malvaceae Mangiaceae Manicariaceae Ma

paniaceae Marantaceae Marathraceae MarcgraviaceaeMartyniaceae Mastixiaceae

Matricariaceae Maundiaceae Mayacaceae Medeolaceae Medusagynaceae Peridiscac

eaeMelaleucaceae Melampyraceae Melanophyllaceae Melanthiaceae Melastomatac

eae Meliaceae Melianthaceae MelicaceaeMeliosmaceae Melittidaceae Melochiacea

e Memecylaceae Mendonciaceae Menispermaceae Menthaceae MenyanthaceaeMe

nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

47

Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

Nypaceae Nyssaceae

Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

ceae Papilionaceae Pappophoraceae PapyraceaeParacryphiaceae Parasitaxaceae Par

ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

48

hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

49

cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

50

ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 7: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

7

The major endosymbiotic events that characterize the clade of which flowering

plants are a part (and gave rise to chloroplasts and mitochondria) are very ancient and

cause no problems for the student of multicellular organisms However lateral gene

transfer has been detected in a number of situations between quite unrelated

organisms (eg Bergthorsson et al 2003 - Amborella and liverworts see also below but

cf Goremykin et al 2009) and it may be particularly common in mitochondria (Sanchez-

Puerta et al 2008) Here too there are no major problems providing one is careful

such transfers do however raise all sorts of interesting biological questions (see

Richardson amp Palmer 2007 for a summary) Although there is increasing evidence for the

importance of genome duplications - hybridization is one cause of this - at various times

during the evolution of seed plants and of palaeopolyploidy events within eg the

Lauraceae and Magnoliaceae clades (Soltis et al 2009 for a summary see eg

the Characters page for further discussion) these too do not currently seem to pose

problems for the adoption of monophyly as the sine qua non of groups to be recognised

formally in this phylogenetic classification but it can make detecting orthologous genes

difficult

The accessory principles of Backlund and Bremer (1998) should be used in

combination Thus keeping the monogenericPlatanaceae separate from its sister

taxon Proteaceae is justifiable Both are much-used names that signal well supported

well defined and easily recognisable groups that have long been recognised as distinct

have several synapomorphies and do indeed look very unlike each other Combining

the two would yield a clade with few obvious apomorphies not to mention the fact

that Nelumbonaceae should by the same logic (it is also monogeneric) also be included

in the expanded family On the other hand it is difficult to justify the continued

recognition of Callitrichaceae or Hippuridaceae monophyletic and distinctive although

they may be If they were recognised several poorly characterised clades would also

have to be carved out ofPlantaginaceae in any classification that aimed to convey a

comprehensive view of the worlds flora The continued recognition

of Valerianaceae and Dipsacaceae also tends to run into this problem (see also Pfeil amp

Crisp 2005 Orthia et al 2005 Albach 2008 etc for useful practical discussions of such

matters) But there are no absolute guidelines If Podostemaceae turn out to be sister

to Hypericaceae (for references here and elsewhere in the Introduction see the

individual families) the subsequent moderate dismemberment of Clusiaceae sl is not

be too high a price to pay for the continued recognition of Podostemaceae Hence the

somewhat provisional recognition of Hypericaceae and Calophyllaceae as well as

Clusiaceae below the families can all be recognized and the name Podostemaceae in

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

8

particular is very well established In a somewhat similar situation if Lemna and its

relatives are a clade sister to most other Araceae should they be recognised as a

separate family Gymnostachys a phenetically fairly distinctive taxon as well as the less

phenetically distinct Orontioideae would have to be recognised as a separate families

(or combined as a single family) too but Araceae in a somewhat restricted sense would

be somewhat more morphologically coherent although not greatly so and not notably

distinct However it is in the very nature of such decisions to be somewhat arbitrary

and unsatisfactory hence the emphasis on consensus classifications here and on

classifications as simply being a means to an end

A useful distinction can be drawn between crown groups and stem groups The

former are monophyletic and include the extant members of a clade and their

immediate common ancestor (see the figure below) The groups characterized in this

site are such groups Thus Proteaceae here are crown group Proteaceae apomorphies

like the single carpel four-merous perianth etc being found in this common ancestor

Stem groups on the other hand include all the members of a lineage immediately after

its split from its sister group and all branches of this lineage In the case of Proteaceae

the stem group would include everything after its split from its sister group

Platanaceae Obviously most of the organisms in the lineage that terminates in crown-

group Proteaceae are unknown only a few being known as fossils and it is not known

exactly where particular apomorphies of crown group Proteaceae evolved along this

lineage

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

9

Indeed there is no reason other than convention or convenience why any group

should not be segregated into several smaller monophyletic groups or merged to

produce a larger unit we can talk about one large thing or about several smaller things

Thinking about aspects of size findings in ethnobiology and cognitive psychology can be

used to suggest that a moderate number - probably fewer than 500 - of families is a

reasonable goal at which to aim and that groupings of taxa throughout any system

should be rather small in size (eg Berlin 1992 Stevens 1994 1997) Major systems such

as those of Linnaeus and Bentham and Hooker were constructed explicitly so as to ease

the burden on the memory (Stevens 1997 2002 see also Scharf 2007) the latter in

particular ensuring that all groups in their classification were relatively small often

containing three to eight immediately subordinate taxa - but by no means all their

groups were formally named Along the same lines Burtt (1977b) suggested that the

number of names at any rank should be at most one third those at the immediately

lower rank - and monotypic taxa might not need a formal name Consistent with such

ideas a fairly broad view of families and orders is taken here whenever the constraints

of monophyly and other criteria used when constructing classifications (see above)

permit The APG system is thus a convention consistent with ideas such as those

expressed by Backlund and Bremer (1998) for similar conventions see eg the Grass

Phylogeny Working Group (2001) and for fungi Hibbett et al (2007)

There are other views Thus Takhtajan (1997) has suggested that smaller families

are more natural This is incorrect Monophyletic groups that include fewer taxa -

Takhtajans smaller families - do not necessarily have more apomorphies than larger

groups even if members of smaller groups will always have more features in general in

common than the larger group that includes them That is they will have their

apomorphies their unique features as well as progressively more plesiomorphies

features found both in the small groups and in the larger clades of which they are a part

having more features in common is indeed one common meaning of more natural

(Note that the implication of the word natural has long been a group of the kind

[usually unspecified] that I think should be recognised and if other botanists had

different ideas too bad their groups were necessarily not natural [see Bather 1927] It

is thus rarely a helpful word and is not used here) Furthermore if this approach is

adopted we will find a slippery slope ahead By this kind of argument all families should

be very small since their members will have a great deal in common and so will be

most natural However as families (for example) are split the relationships that are

evident between the segregates and that were responsible for their being placed in a

single family in the first place will seem to necessitate the recognition of a new order

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

10

etc as is evident in Takhtajans own work - general taxonomic inflation is the result (see

also comparable suggestions in a cladistic context for Brassicales in particular - Ronse de

Craene amp Haston 2006) Such splitting is also questionable when teaching and learning

families since the student needs to understand the system as a whole However for

some genera removed from the families that until now have included them the

phenetic-classificatory-phylogenetic structure in their new home may mandate the

recognition of small families On the other hand Takhtajans suggestion that narrowly

defined families are more useful for phylogenetic studies may be true Indeed I have

more than once regretted prematurely combining groups whether species (in the

context of monographic work) or families (in the course of preparing these notes) but

this is largely a separate issue

Van Steenis (1978) Philipson (1987b) and others have rightly questioned the value

of splitting a group when ideas of the relationships of its constituent members have not

changed - that is very good reasons have to be provided for splitting a family if the

genera within it remain part of the same clade rather than belonging to another clade

Thus APG (2003) broadened the circumscription of Malvaceae because of the

parapolyphyly of some of the families that had historically been associated with it (Judd

amp Manchester 1997 Alverson et al 1999 Bayer et al 1999) These families particularly

Tiliaceae and Sterculiaceae were not at all easy to distinguish their close relationship

(see eg Brown 1814) and overall similarity had long been conceded and to some

workers at least their combination has come as something of a relief Although most of

the larger clades within Malvaceae sl remain difficult to distinguish even with flowers

Cheek (2007) opts for a wholesale and novel dismemberment into ten families

however the very good reasons for doing this are wanting

The same principles are of course applicable when it comes to dividing genera little

other than a headache is gained by splitting genera such

as Drosera and Gnetum (Doweld 2000) as has recently been proposed Thus if an

established genus divides into two (or more) clades this is not a signal for recognising

two groups at the same level - so here it could be argued that the dismemberment

of Pterostylis (Jones amp Clements 2002b) was somewhat unfortunate Along the same

lines if a newly-discovered taxon is sister to an existing named taxon - say a genus - this

does not necessitate the description of a separate genus for the newly described species

(cf Davis 2002) (Of course some [eg Thorne 1976] have suggested that the sizes of

gaps between groups at the same rank should be similar but any principle like this is

inherently flawed since morphological gaps are more unstable than phylogenetic

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

11

relationships and applying it across all flowering plants would both be difficult and cause

substantial changes - see also below) Humphreys and Linder (2009) provide a well-

documented survey of generic concepts in plants which the reader should consult they

note that generic limits (broad versus narrow) have oscillated historically and that

currently larger genera tend to be recognised because studies tend to be on a broader

scale than in the past

Note that invoking similarity or difference - whether qualified (considerable

similarities substantial differences) or not - in a cladistic context as justification for

combining or splitting taxa is not a particularly strong argument (see eg

CardiopteridaceaeStemonuraceae - Karingrehed 2002c) Similarity and difference can

neither be defined precisely since what may seem to be substantial similarities to me

may not to the next person nor are they likely to be stable in the face of our changing

knowledge of morphology and what might be synapomorphies

I might have prefered to merge some families recognised here or split others but

by and large I do not think my own preferences matter very much - and I take the same

position with regards to comparable preferences expressed by others Indeed the

bottom line is that in flagged hierarchies of the kind used here the limits of any

monophyletic unit generally taught and discussed particularly other than species can

be established only by convention and consensus (eg Stevens 2002 2006a Entwisle amp

Weston 2005) This is rather different from the reasonable observation that the

phylogeny itself cannot be achieved by consensus (Thorne 1976) for in most cases there

is a fact of the matter when it comes to relationships Given the increasing support for

the outlines of angiosperm phylogeny a stable consensus classification based on this

phylogeny seems attainable Indeed in addition to providing current ideas of

relationships of seed plants in a synthesised form this site is part of an attempt to build

such a consensus about the circumscription of taxa (see APG 1999 2003 2009 Grass

Phylogeny Working Group 2001 Mabberley 2008) Reaching such a consensus is vital

since what we know of angiosperm phylogeny allows a very large number of

classifications to be based on it and as we find out more the number of possible

classifications increases greatly Unfortunately however nature does not dictate what

the classification should be All classifications are constructed by humans to

communicate particular aspects of groups and relationships Our goals as systematists

are surely to produce robust hypotheses of relationships to understand the evolution of

morphology and the like - but not to argue ad nauseam whether something should be

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

12

a family or a subfamily That way surely lies madness and worse the discredit of our

discipline

There are similar issues whatever naming system is used Thus in phylogenetic

naming (Baum et al 1998 for an example but cf Baum et al 2004 for the PhyloCode

see Cantino amp De Queiroz 2006) an unflagged hierarchy is used in which any

terminations of names used are uninformative about the relative position of taxa If one

adopts the principle of phylogenetic naming one indeed does not have to worry about

the nomenclatural consequences caused by lumping or splitting any well-supported

clade can be named without affecting the name of more or less inclusive clades

Unfortunately unflagged hierarchies have very serious deficiences as communication

devices because they lack one aspect essential in language biological or otherwise -

they contain no intrinsic information about the relationships of the group in question to

others (eg Pfeil amp Crisp 2005 Stevens 2006a) Recent suggestions for using prefixes like

Apo- and Pan- to PhyloCode names will however allow limited information of this

kind to be conveyed but only as it pertains to individual branches and current

proposals do not even mandate that the prefixes be employed consistently In any

event such proposals simply prevent the potential tripling of the number of quite

different names used to describe different aspects of a phylogenetic tree over those

used to name monophyletic groups pure and simple In general where n is the number

of extant species in a group the number of clades in such a group = n-1 (Species will

also need names too for their names see Dayrat et al 2008) Importantly here too

consensus over the clade names commonly learned by students and used in herbaria is

needed otherwise communication will be impeded the names themselves will provide

no guidelines as to which should be chosen The situation is of course more complicated

than this Terminations that convey ideas of rank in a phylogenetic classification can also

be used in phylocode names - however there they will carry no implications of rank

How they will be used is another matter of course

Of course there are other philosophies of classification and some still prefer

evolutionary classifications There classificatory principles differ substantially from those

followed here eg the recognition of paraphyletic taxa may be permitted however

detailed reasons for prefering the taxa that are recognised are rarely given although

nature and natural groups are often mentioned (cf Stuessy amp Koumlnig 2008) For

summaries of commonly used systems see Brummitt (1992) and Mabberley (2008)

new evolutionary systems appear every year or so However even those who allow or

promote the recognition of paraphyletic groups (eg Grant 2003 Thorne 2007

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

13

Heywood et al 2007) may find it of interest to examine a system recognizing only

monophyletic groups it provides a rather different understanding of evolution

Evolutionary classifications in general try and combine phylogeny and

morphological gaps although that is no easy thing to do - it is akin to combining chalk

and cheese (for an attempt to make this impossible task seem to be more objective see

Stuessy amp Koumlnig 2008) However there is a principle from evolutionary classification

that is relevant and I think quite useful here The size of the gap between two groups

tends to be inversely proportional to the sizes of the groups involved (Davis amp Heywood

1963) One can imagine a situation in which a large group is formally divided even

although the distinguishing characters of the two are weak whereas a smaller group

similarly divisable will be left intact

To summarize If hypotheses of phylogeny remain stable we should be able to base

a stable classification on that phylogeny and then get on with our work that is testing

the phylogenies we have elucidating phylogenies in areas where relationships are

unclear studying the evolution of morphology describing species etc In this context

the spread of the Angiosperm Phylogeny Group system (see below) and its widespread

utilisation in technical literature also floras (eg van der Meijden 2005) textbooks (eg

Simpson 2006 Judd et al 2007 [third edition]) dictionaries (Mabberley 2008) more

popular literature (eg Souza amp Lorenzi 2005 Spears 2006) and as an outline for a new

herbarium sequence (Haston et al 2007) is gratifying The posibility that one might be

able to develop a stable phylogeny-based classification of families and in particular

orders represents a dramatic turn-around from the pessimistic attitude about such

higher-level groupings expressed by Davis and Heywood (1963) and Thorne (1976) the

latter even suggesting that we should bury forever the metaphor of the phylogenetic

tree as highly unrealistic (ibid p 56) Returning to Godfray and Knapps (2004) users

of classifications who want a stable informative and accessible classification that

enables easy identification - unfortunately they want cake with everything and cannot

get it - these pages attempt to satisfy as many of their needs as possible but phylogeny

and monophyly are the primary shapers of the APG classification

On this classification in particular

Here I very largely follow the Angiosperm Phylogeny Group classification (APG

2003) Any differences are not to be interpreted as differences in principle simply that

new phylogenies continue to be published and here I attempt to provide an overview

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

14

of current ideas of higher-level relationships of seed plants The Angiosperm Phylogeny

Group classification is based on well-supported relationships evident in the numerous

molecular studies that began to appear in the late 1980s (see APG 1999 for the

principles underlying the classification) but recent phylogenetic work does not

contradict the major outlines of the trees used by APG II (2003) or even those in APG

I (APG 1999) The APG classification has for the most part been conservative and I

am similarly conservative here For the names and authorities I follow APG III (2009)

although the names of the authors may not always be correct these being in something

of a state of flux for the superordinal names used see Chase and Reveal (2009)

although classification at this level here is not exhaustive - only clades with several

orders have superordinal names

Some changes in our ideas of relationships and hence in the clades we talk about

are particularly likely in parts of Lamiales Caryophyllales and Malpighiales for example

although this will not affect the orders themselves Thus some changes to clade

circumscriptions are to be expected although I do not expect them to be substantial

but changes are neither a defect of cladistics nor a necessary consequence of the use of

molecular data Clades are hypotheses of relationships and as hypotheses they may be

overturned However work since APG I has not suggested other than minor changes

in the compositions of the orders even if the odd genus or even family is turning out to

be seriously misplaced - recent examples are Hydatellaceae (from monocots-Poales to

Nymphaeales Saarela et al 2007) Guamatelaceae (from Rosales-Rosaceae to

Crossosomatales Oh amp Potter 2006) and Perrottetia and Bhesa (from Celastrales-

Celastraceae to Huerteales and Malpighiales respectively Zhang amp Simmons 2006) The

main changes have been clarification of the relationhips of individual families or groups

of families that were of uncertain position eg of Chloranthaceae (Moore et al 2007)

Ceratophyllaceae (Jansen et al 2007) and unplaced asterid II families (Winkworth et al

2008a) As mentioned above phylogenies in the area of main interest in these pages are

overwhelmingly tree-like Although there is evidence for acquisition of host plant genes

by parasites (see below) even wider but inexplicable transfer of mitochochondrial

genes in plants like Amborella (Bergthorsson et al 2004 cf Goremykin et al 2009a)

and even transfers of nuclear genes (Vallenback et al 2008) these are the exception

rather than the rule

In cases where the Angiosperm Phylogeny Group II allowed alternatives as to the

limits of families - Papaveraceae in the broad sense or Papaveraceae plus

Pteridophyllaceae plus Fumariaceae Proteaceae in the broad sense or Proteaceae plus

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

15

Platanaceae - the choices made here follow common usage eg as in textbooks like

Judd et al (2007) and Simpson (2006) and particularly in the new edition of

Mabberleys The Plant Book (Mabberley 2008) This latter is itself an attempt to reflect a

consensus the result of taking the opinions of botanists at several meetings A largely

similar consensus is reflected in the APG III classification (APG 2009) where you will

see that alternative classifications have been dispensed with For many people the

existence of such alternative classifications will simply confuse so agreement over

which groupings to use when alternatives are permitted and then dispensing with the

alternatives that are not commonly used seems reasonable I am making the few

changes that are needed to bring the classification here in line with these others but

only gradually since there is not enough time in each day

As already mentioned higher-level relationships in general and the composition of

orders in particular have always presented something of a challenge to systematists

(eg Davis amp Heywood 1963) That being said the composition of some clades like

Apiales Crossosomatales and Pandanales is decidedly unexpected However it is

interesting to see that these higher level clades are generally accepted even in works

with different classificatory philosophies (eg Heywood et al 2007) For clades like

Malpighiales many of the family groupings within Asparagales etc attempts to find

distinctive characters have largely failed (but see Endress amp Matthews 2006a also

the Apomorphies page here) Interestingly as with families some groupings suggested

by molecular studies are supported by morphological andor chemical characters that

have long been known sometimes for over a hundred years the relationship between

Pittosporaceae and ApiaceaeAraliaceae are a case in point (Hegnauer 1969b and

references) As our knowledge of morphology and chemistry improves we can hope for

improvements in the characterisations of clades at all levels

In many cases the new family limits of the Angiosperm Phylogeny Group (see

APG 1999 2003) are not really controversial although changes from the limits

commonly accepted only a decade ago are sometimes quite dramatic (eg Wagenitz

1997) Thus the split of the old Saxifragaceae sl is necessitated by its extreme

polyphyly as also with Icacinaceaesl and Cornaceae sl However the limits and

relationships of such groups had long been considered to be unsatisfactory but until

recently there had been no real reason to prefer one proposed arrangement over

another It is generally accepted that the limits of Lamiaceae and Verbenaceae have to

be redrawn although there is as yet no compelling evidence that the redrawn taxa are

not sister taxa (there is no evidence that they are) But whatever their relationships the

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

16

content of the clades has changed considerably and incidentally they are now easier to

identify than before the decision to recognise two families is not difficult The same is

true for Salicaceae and Achariaceae (Malpighiales) two previously small families that

have received the bulk of the old Flacourtiaceae Clade and hence taxon limits remain

difficult around Theaceae (Ericalessee alsoSladeniaceae Pentaphylacaceae)

and Euphorbiaceae (Malpighiales see

also Phyllanthaceae Picrodendraceae Putranjivaceae etc) Here current groupings

may not represent quite such dramatic changes in our understanding of relationships

Indeed parts of the old Euphorbiaceae that were separated may yet go back together

particularly Phyllanthaceae and Picrodendraceae (see Wurdack et al 2004 Davis et al

2005) even if Theaceae should indeed be dismembered (Geuten et al 2004) However

given that there is no molecular evidence that currently warrants combining all the

segregates of Euphorbiaceae and even if some do come together the clades the

families represent suggest novel groupings not recognised in current classifications

Furthermore Rafflesiaceae appear to be embedded within Euphorbiaceae s str (Davis

et al 2007) so the family is divided Of course maintaining even a moderately narrowly-

circumscribed Euphorbiaceae would entail reducing the iconic Rafflesiaceae to

synonymy Relationships in core Caryophyllales especially

around Phytolaccaceae andNyctaginaceae and also Portulacaceae are incompletely

understood and refashioning of taxon limits will doubtless be needed as cladistic

relationships become apparent (eg see Nyffeler 2007 Ogburn 2007a b) Some

groupings in the old Icacinaceaeand Olacaceae also remain unclear

The discovery of the relationships of parasitic and aquatic groups have presented a

particular challenge to systematists Morphologically some of these plants are so highly

modified that interpretation of the plant body in conventional terms is difficult or even

impossible Thus parasitic groups (broad-leaved angiosperms only) such as Rafflesiaceae

are hard to place since both the vegetative body and the flowers are changed almost

beyond recognition (flowers of rafflesiaceae are a good example of gigantism - Davis et

al 2007) Furthermore plastid gene sequences may be difficult or impossible to obtain

the chloroplast DNA in particular being highly degraded and the rate of molecular

change in general may be high so the problem of long-branch attraction is serious (eg

Duff amp Nickrent 1997 Nickrent et al 1998 Caddick et al 2002a G Petersen et al

2006b) Echlorophyllous sapromycoheterotrophic taxa mostly known from monocots

present similar problems However progress is being made placements for

Apodanthaceae Rafflesiaceae Mitrastemonaceae Cytinaceae and Cynomoriaceae have

recently been suggested (Barkman et al 2004 2007 Davis amp Wurdack 2004 Nickrent et

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

17

al 2004 Davis et al 2007) it seems likely that Burmanniaceae sl are polyphyletic but

both parts are in Dioscoreales (Merckx et al 2006) and relationships within the largely

hemiparasitic Santalales are also gradually being clarified (Maleacutecot 2002) The inclusion

of parasitic taxa in general molecular analyses can cause conniptions (eg Nickrent et al

2004 Davis et al 2004 Chase et al 2006 G Petersen et al 2006b) and there can be

horizontal transmission of genes (eg Davis amp Wurdack 2005 Vitaceae to Rafflesiaceae

Barkman et al 2007 the mitochondrial atp1 gene commonly moves)

Similarly in water plants neither vessels in particular nor much xylem in general is

needed leaves are highly modified and water-mediated pollination if adopted may

well be associated with major changes in floral morphology Here too recent molecular

studies suggest that aquatic groups with hitherto problematic relationships may find

homes Thus Podostemaceae are close to Clusiaceae and Calophyllaceae and are sister

to Hypericaceae (Malpighiales Kita amp Kato 2001) Hydatellaceae which used to be in

Poales are part of Nymphaeales (Saarela et al 2007) and Hydrostachyaceae may be

close to Hydrangeaceae (Cornales Xiang et al 2002) or perhaps should be placed in

Lamiales note that in the first two cases in particular there are morphological and

chemical features that support such a move If Podostemaceae are indeed close to

Clusiaceae I look forward to seeing hypotheses to explain how the dramatic changes in

the vegetative body that have made Podostemaceae so problematic for generations of

systematists took place That conventional wisdom has trouble in understanding or

explaining how the morphologies of groups like Clusiaceae and Podostemaceae can be

related is largely a problem with conventional wisdom Ceratophyllaceae seem to be

finding a position as sister to eudicots (see Moore et al 2007) but their morphology is

so derived that there is no morphological evidence of which I am aware for this

relationship There are also similar although less extreme morphological problems with

many plants pollinated by wind

In such situations especially with some water plants mycoheterotrophs and

parasites a variety of characters may be affected and particularly under such

circumstances groups have tended to be formed using a few or even only a single

character that seems to provide evidence of relationships Add to this the tendency to

weight some characters particularly strongly on a priori grounds and the result was

either the recognition of conglomerate taxa such as Amentiferae which now appear to

be highly polyphyletic or the segregation of families like Plantaginaceae s str (now

much expanded) and Leitneriaceae (now in Simaroubaceae) - all of these are more or

less wind pollinated

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

18

Even although relationships of many parasitic and aquatic groups are being

clarified and at least some of the proposed phylogenetic positions seem to be well

supported it is indeed often still very difficult to understand how such groups have

evolved from their more morphologically conventional relatives However we have had

all sorts of preconceptions as to how characters should evolve Thus we used to

assume that features like highly scalariform vessel perforation plates or the complete

absence of vessels or a flower with an androecium that had many stamens a superior

ovary or separate petals were necessarily plesiomorphic or primitive and conversely

a simple perforation plate and the presence of vessels an androecium with few

stamens inferior ovary or petals that were connnate were almost necessarily

apomorphic or advanced (but cf eg Stebbins 1951) Such assumptions are incorrect

(eg Soltis et al 2005b) Carpels may become secondarily free carpels may fail to close

the seeds then developing outside the confines of the carpel as in some Aspagaraceae-

Nolinoideae Violaceae Berberidaceae Malvaceae-Sterculioideae etc) in Peliosanthes

teta perhaps the only species in Peliosanthes(Asparagaceae-Nolinoideae) the ovary

varies from superior to inferior (Jessop 1976 species limits here need close

investigation see also Kuzoff et al 2001 and Soltis amp Hufford 2002 Saxifragaceae

Apiales etc) many-seeded carpels can evolve from few-seeded carpels

(Razafimandimbison et al 2008) monoecy may be derived from dioecy (Schaefer amp

Renner 2010 and references) Classic studies such as those by Babcock (eg 1947)

on Crepis that assumed that evolution - in this case of the karyotype in particular - was

unidirectional have needed comprehensive re-evaluation (Enke amp Gemeinholzer 2008)

Most if not all characters have reversed andor evolved in parallel as is clear in the

discussion of ovary position here especially in Poales Saxifragales and Asparagales and

also in comments on such morphologically distinctive taxa (when compared with their

immediate relatives) such as Menyanthaceae and especially Pittosporaceae Parallel

evolution may occur even at the level of amino acid substitution as in the independent

acquisition of the phosphoenolpyruvate carboxylase (pepC) gene in C4 photosynthesis in

grasses (Christin et al 2007b see also Blaumlsing et al 2000)

Thinking of how such characters have been used in the past in classifications at the

generic level the results have been similar to those just discussed above for parasitic

plants and others Relying too much on animal pollination syndromes to mark generic

boundaries has all too often led to taxa that are highly unsatisfactory phylogenetically

(see eg Acanthaceae Bignoniaceae Campanulaceae Ericaceae Lamiaceae

Orchidaceae) and over-reliance on characters of fruit and seed (see eg Brassicaceae

and Apiaceae) has also led to unsatisfactory generic limits Again the more general

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

19

problem is the use of one or a very few characters that have been weighted a priori to

structure classifications (see also Garciacutea et al 2009) although it is perhaps not

surprising that such an approach sometimes results in the delineation of taxa that have

indeed turned out to be quite distinct (cf eg Burtt Astragalus versus Oxytropis)

Taxa recognized here are monophyletic yet many are polythetic at the

morphological level that is they lack unique features characterizing (defining) all and

only members of that family they can be recognised phenetically only by the unique

combinations of characters that they posess This is the result of evolution any

synapomorphy characterizing a taxon may be lost or modified beyond easy recognition

in some of its members or the synapomorphy may appear to be identical to a feature

that has evolved in parallel in a quite unrelated plant That plant groups are polythetic is

almost as much a feature of monophyletic taxa as the paraphyletic taxa common in

evolutionary classifications Some families now include substantial variation as

phenetically distinct derived groups are placed in their proper phylogenetic position -

examples are Ericaceae which include the erstwhile Empetraceae a wind-pollinated

group that is florally very different and the overwhelmingly large-flowered and animal-

pollinated Plantaginaceae which now include several derived small-flowered aquatic

and wind-pollinated groups that were previously placed in separate families

Thus some families as delimited here may not be easy to recognize However

remember that detecting relationships - use whatever characters you can even if they

are not obvious - and naming a plant - focus on easy-to-see characters that may not

reflect relationships - are quite different problems Taxa although natural may not be

readily recognizable indeed it was in exactly this context that Lamarck worked out the

basic principles of writing dichotomous keys in 1778 (see Scharf 2007 for keys and the

like) Of course Lamarcks idea of nature was very different from ours - he thought that

there was some kind of continuum of form on which living organisms were to be

situated with no real gaps anywhere - but this meant that his genera (for example)

might well not be sharply distinct from each other even if each were part of the real

continuum that was life Lamarcks separation of the establishment of relationships and

groupings in a system and identification of the plants in that system have been widely

accepted (but cf eg Godfray amp Knapp 2004 many of the contributors to Heywood et

al 2007) Perhaps the best way of identifying plants at the family level is by well-made

multiple access keys as in Watson and Dallwitz (1992a onwards family limits there may

differ substantially from those adopted here) Multiple access keys free users from the

constraints of dichotomous keys in which particular characters are needed at each step

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

20

of the identification process before there can be further progress Instead those

characters that happen to be evident on a specimen can be used in whatever order is

convenient when linked to illustrations glossaries etc their power is enormous (see

Dallwitz et al 2000 [2006] for the principles underlying their construction and use)

Nevertheless dichotomous keys such as those of Hutchinson (1973) and Franz Thonner

(Geesink et al 1981) have their uses Of course taxa coming out adjacent in keys may

well not be at all related

(When identifying large numbers of plants even more efficient than either style of

identification and certainly lots more fun is sight identification Unless you have a

photographic memory you have to build up your knowledge of comparative plant

morphology - on which the ability to make accurate identifications depends - by

repeated observation When faced with an unknown plant I always look for leaf teeth

and stipules distinctive hair types especially stellate and T-shaped hairs glands and

punctations of any sort the presence of latex or other exudate and check leaf insertion

smelling crushed leaves can also be helpful In this context nodal anatomy can usually

be checked using a razor and a hand lens or even simply carefully examining leaf scars

The short paragraphs added after most families may help in confirming familial

identifications)

For the record and for the little that it is worth there are 4 orders and 13 families

of gymnosperms characterised on these pages and together they include some 82

genera and 947 species For angiosperms comparable figures are 56 orders 445

families 13208 genera and 261750 species (of which monocots include 11 orders 89

families 2759 genera and 52760 species) Note however that higher mathematics

was never my strong point and anyway these are pretty meaningless figures even for

species which many (but not all) might concede smacked slightly more of reality than

other taxa estimates range as high as 422000 (Govaerts 2001) Furthermore numbers

of genera and species change daily Nevertheless as emphasized here families are

useful in teaching we as a community can ensure that their limits remain largely stable

and by concentrating on relatively few of them one can gain some familiarity with much

of the worlds flora For summaries of the sizes of orders and families and of the general

arrangement followed in these pages see also the Statistics page itself

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

21

A PHYLOGENETIC TREE OF SEED PLANTS (THE MAIN TREE)

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

22

The Main Tree above is a conservative summary of well supported relationships

between the major clades of extant seed plants Reading - interpreting and

understanding - such a phylogenetic tree is not necessarily easy so reading a tree may

require a bit of practice OHara (eg 1988 1992) provides an accessible introduction to

thinking and talking about history in the context of looking at phylogenetic trees More

recently Krell and Cranston (2004) Crisp and Cook (2005) and others have emphasized

how careful one must be when interpreting and talking about evolution in ladderized

trees in particular and phylogenetic trees in general Gregory (2008) provides a

particularly detailed discussion of this problem In ladderized trees the smaller (in terms

of numbers of terminals) sister taxon is consistently shown on the same side at every

node thus the trees tend to be pectinate like the teeth of a comb however the

horizontal axis that is as a result so evident has no polarity or particular direction The

use of the adjective basal when discussing phylogenetic trees is especially dangerous

(see also Wojciechowski et al 2004 D Soltis et al 2005b) When I use the term and the

context is not otherwise clear I am referring to the pectinations at the base of a

ladderised tree Remember that when talking about sister taxa one can never be basal

to the other or older or younger although members of one can be more derived - in the

sense of having more apomorphies - than the other (but even then one has to be

careful) The words primitive and advanced and lower and higher should

practically never be used especially when talking about taxa Amborellaceae and

Pinaceae are sister to all other flowering plants and Pinales and are basal to the crown

groups of all other flowering plantsPinales respectively but that does not mean they

are thereby primitive Pinaceae in particular have numerous apomorphies The word

plesiomorphic is far less loaded than primitive and can be used to talk about

individual characters

Most trees here have been more or less ladderized as is the one above The

ladderization may be imperfect for example you can see that on the Main Tree

asterids with ten orders follow rosids and relatives which have seventeen although

this is in part because of the currently unresolved nature of relationships between the

main clades of core eudicots (this is likely to change soon) The Main Tree could have

been drawn with Amborella Acorus or a host of other taxa at the far right without

offending any relationships phylogenetic trees are like mobiles the only fixed points

being the nodes Despite the problems just mentioned pectination interpreted

carefully has its value and it is not an arbitrary process As one reads the terminals of a

pectinate tree from left to right adjacent terminals will be separated by apomorphies

that will cumulate in an appropriate fashion When thinking of a book or a herbarium

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

23

sequence (see Haston et al 2007) this is of some value Nymphaeales and

Austrobaileyales are here adjacent on the tree but they could be separated by

hundreds of families in the sequence merely by rotating only the node from which

Nymphaeales and their sister group (all other angiosperms minus about 75 species)

arise If adjacent in a book or herbarium or as in these summary pages then it is

relatively easy to relate their characters both apomorphic and plesiomorphic but if

separated by hundreds of pages or two floors in a large building then it is less easy to

get anything from the sequence (Since all orders in the pages of the main part of this

site are preceded by the apomorphies of all nodes immediately below them in the seed

plant phylogeny and because of the linkages that have been built in to the site this

problem is much less serious there there is no sequence) So the particular sequence of

all seed plants that is based on a phylogenetic classification but is to be used in herbaria

or any other place where taxa have to be arranged linearly as in this part of APweb

will be one that maximizes the number of taxa that are both successive branches of the

tree and placed successively in the sequence Since specimens are generally filed under

families the outline of a new family sequence for arranging herbaria and books can now

be suggested (Haston et al 2007) and such a standardized sequence will further help

teaching and learning about plants

ON THINKING ABOUT APOMORPHIES

Identifying apomorphies is important because understanding the evolution of

morphology in the broad sense - ie including anatomy chemistry etc - in the context

of diversification is one of our major goals For identifying apomorphies several

preconditions must be met One needs to have an accurate well supported phylogeny

one has to have examined the right taxa both from the point of view of morphology and

molecules one has to have coded the characters right (ie delimited states

appropriately) and one has to use an appropriate model of evolution when fitting the

variation to the tree (see eg Omland 1999 Stevens 2006b) If the distributions of

apomorphies below are compared with those in other studies (eg Turgeon et al 2001

Bremer et al 2001 Endress 2001 Albach et al 2001a Judd amp Olmstead 2004 D Soltis

et al 2005b Zhang et al 2006 Judd et al 2007) the reader may find a number of

differences Although I have integrated such studies as far as is possible there are five

reasons that link to the preconditions just mentioned why there may be differences

where features are placed on trees

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

24

1 I may not have found all the information about a particular character there may be

disagreement over its interpretation or I have added information

2 The sampling of nearly all molecular studies is very incomplete (see Salisbury amp Kim

2001 for problems caused by sampling) indeed it is commonplace to decry the

incompleteness of molecular sampling But not only is the sampling in molecular studies

often less than we might wish that of the morphological and chemical characters whose

evolution we are interested in understanding is also often very poor So for many

anatomical chemical and embryological characters that are confidently said to

characterise families and other groups we all too often have no idea if those characters

are applicable to the whole clade or just to a subgroup within that clade However as

our sampling improves we can locate changes on the tree more precisely if in

unexpected positions Thus is is sometimes suggested that Ericaceae have ellagic acid

(Soltis et al 2005b) in fact the little evidence we have indicates that only a small clade

of some 80 species has ellagic acid and the rest of the family - itself alone about one

third of all Ericales - does not Similarly Ericaceae are noted for their

ectendomycorrhizal associations but Enkianthus sister to all other Ericaceae

apparently lacks these (Abe 2005) Thus these mycorrhizae and their associated

distinctive hair roots are unlikely to be apomorphies of Ericaceae - however it appears

that other families near Ericaceae may also have ectendomycorrhizae (see Asai 1934)

3 I am fitting characters to a very conservative tree with many polytomies although the

nodes that are utilised are for the most part strongly supported polytomies make the

optimisation of characters that is the assigment of character state change to a

particular node on the tree notably difficult (eg Madison amp Madison 2002) In nearly

all studies of the evolution of characters distributions of characters are optimised on a

more or less fully resolved tree and the construction of supertrees may yield yet more

detailed hypotheses of relationships (for literature on supertrees see Cotton amp

Wilkinson 2007 2008) Of course some nodes on such fully resolved trees andor

supertrees may have little support and optimisations of characters on such trees may

carry correspondingly little conviction Even parts of some of the trees used here have

poor support eg relationships within aquatic Alismatales etc although I indicate

when this is the case

4 Exactly how one goes about optimising a character on a tree is critically important

Even using simple parsimony optimisations (ACCTRAN or DELTRAN ACCelerated

TRANsitions or DELayed TRANsitions) the position of synapomorphies on trees - and

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

25

hence our ideas of evolution - may differ and this is still more true if one uses maximum

likelihood or Bayesian methods and makes apparently reasonable suggestions about

weighting gains over losses (eg Donoghue amp Ackerley 1996 Cunningham et al 1998

Omland 1997 1999 Ree amp Donoghue 1999 Polly 2001 Webster amp Purvis 2001

Ronquist 2004 Crisp amp Cook 2005) Sannier et al (2007) show how in in Arecaceae that

where on a phylogenetic tree one might peg changes in microsporogenesis will depend

on the methods one uses to do this and Pedersen et al (2007) discuss the sometimes

very substantial effect of node support on the posterior probabilities of ancestral

character states Here I use parsimony optimization not always as explicit as it might be

but I have often indicated where there are particularly important uncertainties as to the

positions of particular character changes on the tree

5 Finally although I have paid quite a lot of attention to the delimitation of the

character states that make up all the characterizations I have not spent enough time on

this critical operation If we are interested in understanding evolution then fitting the

basic variation - not character states - to a tree in principle allows greater flexibility in

understanding morphology in the context of local phylogenies (see also Stevens 2000

Endress 2005c) However many character states used here are delimited globally that

is they are circumscribed in the context of the variation shown by individual characters

across all angiosperms andor in the context of classic ideas of character evolution

Character states often have arbitrary limits and serve best to communicate

information whether they are in fact suitable for either phylogenetic analysis or

understanding evolution are separate issues Studies have rather unsurprisingly

perhaps but importantly shown that dividing the one character into different sets of

states may yield differing ideas of evolution of that character (eg Lamb Frye amp Kron

2003 Hibbett 2004) When looking at trees on which character states are optimised

one should bear in mind the problems surrounding the delimitation of states (eg

Stevens 2000 2006b) and the danger of using pollen or other types - constructs

based on many characters that vary independently but which effectively get lost in

these types thus Blackmore et al (2009) decomposed the pollen types in Asteraceae

into 52 characters

I use Remanes three main criteria of homology or better similarity when

determining the basic similarity of structures on different organisms (see Remane 1952)

These criteria are special properties position and intermediates Special properties

include anatomical or chemical characters gene expression data etc Position refers

to the position of an organ with respect to landmarks on the plant Although plants are

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

26

plants and landmarks are not as fixed as one might like trying to work out the relative

positions of parts is a good way of understanding morphology so long as one does not

try to out-do Procrustes Intermediates include intermediates found on the same

plant or on different plants Intermediates may observed during development or be

adult structures Thus one may find changes during development which suggest what

the nature of a particular structure is and very different stuctures on different plants

may turn out to be similar early in development Or one can simply compare different

taxa and see that structures that appear to be very different when comparing two

different taxa can be linked morphologically by looking at other taxa

Nevertheless the use of these criteria may not yield an unambiguous answer as to

what a structure is even given a solid phylogeny and an improved understanding of

development (see Jaramillo amp Kramer 2007 for a useful discussion) As Endress (2005c)

observed a number of features - position function development shape anatomy

histology gene activity and relationships to other taxa that clearly have petals - can be

used to distinguish a petal (for example) from other floral structures if a petal does not

have one of these features is it thereby not a petal Thus Maturen et al (2005) recently

found that floral organ diversity genes (B and C) were expressed in the large white

inflorescence bracts of Cornus (see also Costa et al 2005) Peney et al (2005) noted that

not all monosulcate pollen grains in monocots have the same developmental pathway

and that as a result such pollen might not have the same ancestral state Reeves and

Olmstead (2003) suggested that the genetic mechanisms causing monosymmetry in

Lamiales and Solanales were different and Serna and Martin (2006) described similar

problems with the development of hairs in Arabidopsis when compared with that of

hairs in Antirrhinum and Solanaceae Indeed as one perhaps might expect delimitation

of states and characters does not necessarily become easier with increasing knowledge

of development etc Thus Buzgo et al (2004) Matthews and Endress (2005) and others

have shown how hard it can be to distinguish between eg prophylls and other floral

structures as their behaviour is studied during the course of the development of the

flower Similarly at what concentration is a particular secondary metabolite deemed to

be present (Waterman (2007)

The validity of the approach used here that of fitting morphological variation to a

largely molecular-based tree may be questioned However I think it rather unlikely that

well-supported molecular branches will be overturned by morphological data Indeed

analyses of morphological data alone do provide support for many of the clades evident

in molecule-only analyses and analyses with morphological and molecular data

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

27

together may lead to increased support for clades (eg Hufford 1992 Nandi et al 1998

[but adding morphological data reduces support for a number of critical clades too]

Doyle amp Endress 2000 however in none of these papers is the use of morphology

without ambiguity) It is unfortunately clear that the use of morphology alone may not

suggest problems in the phylogenetic placement of taxa that later turn out to have been

wrongly included (eg Zhang et al 1992) Thus I have been wary of putting much weight

on clades that have only morphological support but note that I have treated molecule-

based clades with low bootstrap or jacknife support values (esp below 70) or low

posterior probabilies (below 095) likewise Although I may have been mistaken in

placing so much emphasis on molecular data in terms of providing the basic

phylogenetic framework for angiosperms morphological and molecular data are only

very rarely in irreconcilably strong conflict There are indeed a few places where the

conflict seems extreme These include the relative positions of the Monimiaceae and

Hernandiaceae (Laurales) the position of Hanguanaceae (Commelinales [as here] or

Zingiberales) and of Triplostegia (is it in Dipsacaceae or Valerianaceae - see

Dipsacales) Fossils are unlikely to affect the topologies of the trees presented here but

see below for their importance in understanding morphological evolution in general and

the evolution of angiosperms in particular Nevertheless some largely reject the idea

that trees based on molecular data alone can recover phylogenetic relationships

especially when branching points are old and prefer to used trees based on analysis of

morphological data including those taken from fossils (Hilton amp Bateman 2006 Farjon

2007)

All in all however the extent of the congruence between morphological and

molecular data is impressive and heartening and many clades can be characterised

morphologically It seemed in 1998 that there were no unambiguous morphological

synapomorphies for angiosperm orders (K Bremer 2000) and this is still true if by

unambiguous is meant non-homoplasious However many orders and other clades

have synapomorphies even if these may be indistinguishable at least at the current

level of morphological and developmental knowledge from parallel occurences

elsewhere As our morphological knowledge increases so too does the number of

apomorphies

Endress and Matthews (2006a) emphasize the importance of tendencies and

developmental constraints when thinking about characters of clades within the rosids -

many characters or character combinations occur in a rather sporadic fashion within

the clade and apparently notably less frequently outside it There are a number of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

28

examples of tendencies here and in the individual order pages such as the features

enclosed by parentheses in groups above the level of family eg see (cuticular waxes

as aggregated rodlets) for the commelinids and the discussion of the distribution of

polyandry that is flowers with numerous stamens in the asterid I + II groups etc

Tendencies may involve more than single characters When thinking of the

associations of features like integument thickness and vascularization of the integument

(Endress amp Matthews 2006a) one can imagine fairly simple developmental

preconditions being operative As Endress and Matthews (2006a) note it is difficult to

imagine vascular tissue developing in an integument only two cells thick Changes here

would be loosely correlated if morphologically linked However the strongly correlated

changes noted by Givnish et al (2005) are ecologically linked but are presumably

morphologicallydevelopmentally independent When there seem to be characters

evolving more or less together tests can be carried out to see if the changes are

concentrated on certain branches of the tree (eg Maddison 1990 Sanderson 1991

Maddison amp Maddison 2000) Such tests have rarely been carried out at higher levels in

angiosperms

One of the most striking examples of a tendency is the distribution of N-fixation

restricted as it is to a monophyletic group of four clades although it has arisen seven (or

perhaps several more) times independently within the N-fixing clade and several

members of two quite different kinds of bacteria are involved (eg Clawson et al 2004

Elliott et al 2007 Sprent amp James 2007) Other examples of work that bears on the issue

of tendencies include the findings that flowers of polysymmetrical Arabidopsis have

genes like TCP1 that are expressed asymmetrically during early development and TCP1

is a probable orthologue of the well-known CYC gene of Antirrhinum that is involved in

the development of monosymmetric slowers there (Cubas et al 2001 Costa et al 2005

etc) Parallelism might build on this underlying morphologically cryptic monosymmetry

even if details of the genetic mechanisms causing the monosymmetry evident in

particular groups may be different (Reeves amp Olmstead 2003 Cubas 2004 see above)

Similarly there have been several recent suggestions that the capability to synthesise a

particular metabolite may be switched off but not lost and so can sometimes be

reacquired (eg Wink amp Witte 1983 Wink 2003 Liscombe et al 2005 Larsson 2007

Waterman 2007) Hence perhaps the rather spotty distribution of many secondary

metabolites like ellagic acid the indole alkaloid camptothecin iridoids etc when

considered in the context of phylogenies As a non-botanical example - but a rather nice

one - Salwini-Plawen and Mayr (1961) suggested some time ago that there has been

considerable parallelism (40-65 or more independent origins) in the evolution of eyes in

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

29

metazoans However Pax 6 seems to be a master control gene that is involved in eye

formation perhaps throughout all bilateralians (eg Gehring amp Ikeo 1999 Erwin amp

Davidson 2002) which changes the nature of the problem Note however that in

general our invocation of evolutionary tendencies to explain the patterns of

distributions of characters is really an admission that we do not understand those

patterns

As we find out more about variation we will probably find fewer and fewer features

constant throughout a group Most unqualified statements of presence and absence

should properly be qualified as usually present or usually absent if one is thinking of

the characterisations on the order pages as encompassing the total variation within a

clade Thus Pistia alone among monocots as so far known has sieve tube plastids with

starch grains not protein crystals However this does not affect the fact that sieve tube

plastids with cuneate protein crystals and lacking starch grains are an apomorphy for

monocots Furthermore fossils in a number of cases suggest character combinations

unknown in extant taxa as may be seen in the discussions of Fagaceae Platanaceae

Iteaceae Calycanthaceae etc Confusing the issue there may be questions as to where

exactly on the tree a particular fossil is to be placed (eg see Nymphaeaceae

Calycanthaceae Archaefructus etc)

To summarize given our current understandings of both phylogenies and

characters evolution of some characters in which we are interested seems very labile

(see eg D Soltis et al 2005b Endress and Matthews 2006a Stevens 2006b Ekman et

al 2008 for a good recent study) and I have been cautious when talking about character

evolution Much effort must continue to be spent in summarizing characters of well-

established clades at all levels providing features by which they may be recognized and

signaling synapomorphies Remember that (1) the basic morphological anatomical and

chemical knowledge of many critical taxa is woefully incomplete (2) different

assumptions about character evolution may greatly affect the position of

synapomorphies on trees (3) in many cases relationships within and between many

groups are too uncertain at present to worry very much about synapomorphies and

(4) we must be clear about what we do and do not not know As mentioned above I

have indicated in a number of places where there are particularly important

uncertainties as to where characters should be placed on the tree Nevertheless it is a

relatively easy matter to update notes such as these and it can be a simple matter to

incorporate new data on characters that have never before been considered in the

context of a tree Much basic - and unfortunately perhaps unfashionable - work must be

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

30

carried to clarify the distribution of morphological anatomical and chemical characters

for examples of what can to be done see the work of P K Endress and collaborators

(floral morphology and development) and S R Jensen and collaborators (iridoids) But

acquiring information about nodal anatomy cell and tissue distribution nodal anatomy

and the like is not difficult at all and should be generally encouraged

But all this is merely a necesary prelude to the understanding of evolution For this

one needs to know a lot more including dating the phylogeny understanding the

function(s) of characters factoring in relevant aspects of the palaeoenvironment in

which an apomorphy first appeared etc

SUMMARY OF THE SYSTEM

Below is a formalised summary of the relationships within orders of the families of seed

plants There are a few families that are not recognised even as options in APG II and

vice versa as well as a few extra orders The families to be recognised in the most

recent edition of Mabberleys The Plant Book (Mabberley 2008) are also largely

consistent with those below But all differences are trivial and will - I hope - eventually

disappear

Square brackets - [] - enclose clades the plus sign - + - designates sister taxa a

comma - - denotes part of a polytomy and quotation marks - - denotes a

paraphyletic group

SEED PLANTS

GYMNOSPERMS

Cycadales

Cycadaceae + Zamiaceae

Ginkgoales

Ginkgoaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

31

Pinales

Pinaceae [[Araucariaceae [Podocarpaceae + Phyllocladaceae]] [Sciadopityaceae

[Taxaceae + Cupressaceae]]]

Gnetales

Ephedraceae [Gnetaceae + Welwitschiaceae]

ANGIOSPERMSFLOWERING PLANTS

Amborellales

Amborellaceae

Nymphaeales

Hydatellaceae [Cabombaceae + Nymphaeaceae]

Austrobaileyales

Austrobaileyaceae [Schisandraceae + Trimeniaceae]

Chloranthales

Chloranthaceae

MAGNOLIIDS

Magnoliales

Myristicaceae [Magnoliaceae [[Himantandraceae + Degeneriaceae] [Eupomatiaceae +

Annonaceae]]]

Laurales

Calycanthaceae [[Siparunaceae [Gomortegaceae + Atherospermataceae]] [Monimiaceae

[Hernandiaceae + Lauraceae]]]

Canellales

Canellaceae + Winteraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

32

Piperales

[Hydnoraceae + Aristolochiaceae] [Piperaceae + Saururaceae]

MONOCOTS

Acorales

Acoraceae

Alismatales

Araceae [Tofieldiaceae [[Alismataceae [Hydrocharitaceae + Butomaceae]]

[Scheuchzeriaceae [Aponogetonaceae [Juncaginaceae [Maundiaceae [[Posidoniaceae

[Ruppiaceae + Cymodoceaceae]] [Zosteraceae + Potamogetonaceae]]]]]]]

Petrosaviales

Petrosaviaceae

Dioscoreales

Nartheciaceae [[Taccaceae + Thismiaceae] [Burmanniaceae + Dioscoreaceae]]

Pandanales

Velloziaceae Triuridaceae Stemonaceae [Pandanaceae + Cyclanthaceae]

Liliales

Corsiaceae [Campynemataceae [Petermanniaceae [Colchicaceae + Alstroemeriaceae]]

Melanthiaceae [[Philesiaceae + Rhipogonaceae] [Smilacaceae + Liliaceae]]]

Asparagales

Orchidaceae [[Boryaceae [Blandfordiaceae [Lanariaceae [Asteliaceae + Hypoxidaceae]]]]

[[Ixioliriaceae + Tecophilaeaceae] [Doryanthaceae [Iridaceae [Xeronemataceae

[Xanthorrhoeaceae [Alliaceae + Asparagaceae]]]]]]]

COMMELINIDS

Unplaced

Dasypogonaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

33

Arecales

Arecaceae

Poales

[Typhaceae + Bromeliaceae] [Rapateaceae [[Thurniaceae [Juncaceae + Cyperaceae]]

[[Mayacaceae [Eriocaulaceae + Xyridaceae]] [Flagellariaceae [Anarthriaceae

[Centrolepidaceae + Restionaceae]]] [Joinvilleaceae [Ecdeiocoleaceae + Poaceae]]]]]]]

Commelinales

[Commelinaceae + Hanguanaceae] [Philydraceae [Haemodoraceae + Pontederiaceae]]

Zingiberales

Musaceae [Strelitziaceae + Lowiaceae] Heliconiaceae [[Cannaceae + Marantaceae]

[Costaceae + Zingiberaceae]]

Ceratophyllales

Ceratophyllaceae

EUDICOTS

Ranunculales

Eupteleaceae [Papaveraceae [[[Lardizabalaceae + Circaeasteraceae] [Menispermaceae

[Berberidaceae + Ranunculaceae]]]]

Sabiales

Sabiaceae

Proteales

Nelumbonaceae [Platanaceae + Proteaceae]

Trochodendrales

Trochodendraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

34

Buxales

Haptanthaceae [Buxaceae + Didymelaceae]

CORE EUDICOTS

Gunnerales

Gunneraceae + Myrothamnaceae

Dilleniales

Dilleniaceae

Saxifragales

Peridiscaceae [[Paeoniaceae [Altingiaceae [Hamamelidaceae [Cercidiphyllaceae +

Daphniphyllaceae]]]] [[Crassulaceae [Aphanopetalaceae [Tetracarpaeaceae

[Penthoraceae + Haloragaceae]]]] [Iteaceae [Grossulariaceae + Saxifragaceae]]]]

Cynomoriaceae unplaced

Vitales

Vitaceae

ROSIDS

FABIDROSID I

Zygophyllales

Krameriaceae + Zygophyllaceae

Celastrales

Lepidobotryaceae + Celastraceae

Oxalidales

Huaceae [[Connaraceae + Oxalidaceae] [Cunoniaceae [Elaeocarpaceae [Brunelliaceae +

Cephalotaceae]]]]

Malpighiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

35

[Achariaceae [Goupiaceae [Violaceae + Passifloraceae] [Lacistemataceae Salicaceae]]

[Lophopyxidaceae Putranjivaceae] [Ctenolophonaceae [Erythroxylaceae +

Rhizophoraceae]] Linaceae Ixonanthaceae Humiriaceae Irvingiaceae

Centroplacaceae Pandaceae [[Bonnetiaceae + Clusiaceae] [Calophyllaceae

[Hypericaceae + Podostemaceae]]] [Malpighiaceae + Elatinaceae] Ochnaceae

[Peraceae [Rafflesiaceae + Euphorbiaceae]] [Phyllanthaceae + Picrodendraceae]

[Balanopaceae [[Trigoniaceae + Dichapetalaceae] [Chrysobalanaceae +

Euphroniaceae]]] Caryocaraceae

N-FIXING CLADE

Fabales

Quillajaceae [Fabaceae [Polygalaceae + Surianaceae]]

Rosales

Rosaceae [[Barbeyaceae [Elaeagnaceae Dirachmaceae Rhamnaceae]] [Ulmaceae

[Cannabaceae [Moraceae + Urticaceae]]]]

Cucurbitales

Anisophylleaceae [[Corynocarpaceae + Coriariaceae] [Cucurbitaceae [Tetramelaceae

[Datiscaceae + Begoniaceae]]]] Apodanthaceae

Fagales

Nothofagaceae [Fagaceae [[Myricaceae + Juglandaceae] [Casuarinaceae

[Ticodendraceae + Betulaceae]]]]

MALVIDROSID II

pgtGeraniales

Geraniaceae [[Melianthaceae + Francoaceae] [Vivianaceae + Ledocarpaceae]]

Myrtales

Combretaceae [[Onagraceae + Lythraceae] [[Vochysiaceae + Myrtaceae]

[Melastomataceae [Crypteroniaceae [Alzateaceae + Penaeaceae]]]]]

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

36

Crossosomatales

[Staphyleaceae [Guamatelaceae [Crossosomataceae + Stachyuraceae]]] [Aphloiaceae

[Geissolomataceae + Strasburgeriaceae]]

Picramniales

Picramniaceae

Sapindales

Biebersteiniaceae Nitrariaceae [Kirkiaceae [Anacardiaceae + Burseraceae]]

[Sapindaceae [Simaroubaceae Rutaceae Meliaceae]]

Huerteales

Gerradinaceae [Dipentodontaceae + Tapisciaceae]

Malvales

Neuradaceae [Thymelaeaceae [Sphaerosepalaceae Bixaceae [Cistaceae

[Sarcolaenaceae + Dipterocarpaceae]] [Cytinaceae + Muntingiaceae] Malvaceae]]

Brassicales

[Akaniaceae + Tropaeolaceae] [[Moringaceae + Caricaceae] [Setchellanthaceae

[Limnanthaceae [[Koeberliniaceae [Bataceae + Salvadoraceae]] [Emblingiaceae

[Pentadiplandraceae [Gyrostemonaceae Resedaceae Stixis and relatives] Tovariaceae

[Capparaceae [Cleomaceae + Brassicaceae]]]]]]]]

Berberidopsidales

Aextoxicaceae + Berberidopsidaceae

Santalales

Erythropalaceae Olacaceae [[Loranthaceae [Misodendraceae + Schoepfiaceae]]

[Opiliaceae + Santalaceae]] Balanophoraceae unplaced

Caryophyllales

[[Droseraceae [Nepenthaceae [Drosophyllaceae [Ancistrocladaceae +

Dioncophylleaceae]]]]] [[Frankeniaceae + Tamaricaceae] [Polygonaceae +

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

37

Plumbaginaceae]]] [Rhabdodendraceae [Simmondsiaceae [[Asteropeiaceae +

Physenaceae] [[Caryophyllaceae [Achatocarpaceae + Amaranthaceae]]

[Stegnospermataceae [Limeaceae [[Lophocarpaceae [Barbeuiaceae [Aizoaceae

[Sarcobataceae Phytolaccaceae Nyctaginaceae]]]] [Molluginaceae [Halophytaceae

Montiaceae Basellaceae Didiereaceae [Talinaceae [Portulacaceae [Anacampseros etc +

Cactaceae]]]]]]]]]]]]]

ASTERIDS

Cornales

[Cornaceae + Nyssaceae] [Hydrangeaceae + Loasaceae] Hydrostachyaceae [Curtisiaceae

+ Grubbiaceae]

Ericales

[Balsaminaceae [Marcgraviaceae + Tetrameristaceae]] [[Polemoniaceae +

Fouquieraceae] Lecythidaceae [[Sladeniaceae + Pentaphylacaceae] [Sapotaceae

[Ebenaceae [Maesaceae [Theophrastaceae [Myrsinaceae + Primulaceae]]]]]

[Mitrastemonaceae Theaceae [Symplocaceae [Styracaceae + Diapensiaceae]]

[[Sarraceniaceae [Roridulaceae + Actinidiaceae] [Clethraceae [Cyrillaceae +

Ericaceae]]]]]]

LAMIIDASTERID I

Unplaced

Metteniusaceae Oncothecaceae Icacinaceae and some unplaced ex-Icacinaceae

Garryales

Garryaceae + Eucommiaceae

Unplaced

Boraginaceae Vahliaceae

Gentianales

Rubiaceae [Gentianaceae [Loganiaceae [Gelsemiaceae + Apocynaceae]]]

Lamiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

38

Plocospermataceae [Oleaceae [Tetrachondraceae [Calceolariaceae

[Gesneriaceae Peltanthera [Pedaliaceae Carlemanniaceae Martyniaceae Stilbaceae

Plantaginaceae Linderniaceae Bignoniaceae Verbenaceae Lamiaceae Paulowniaceae

Schlegeliaceae Thomandersiaceae Phrymaceae Rehmannia Acanthaceae

Scrophulariaceae Orobanchaceae Byblidaceae Lentibulariaceae]]]]]

Solanales

[Montiniaceae [Sphenocleaceae + Hydroleaceae]] [Convolvulaceae + Solanaceae]

CAMPANULIDASTERID II

Aquifoliales

[[Cardiopteridaceae + Stemonuraceae] [Aquifoliaceae [Helwingiaceae +

Phyllonomaceae]]

Asterales

[Rousseaceae + Campanulaceae] Pentaphragmataceae [[Alseuosmiaceae [Phellinaceae

+ Argophyllaceae]] Stylidiaceae [Menyanthaceae [Goodeniaceae [Calyceraceae +

Asteraceae]]]]

Escalloniales

Escalloniaceae

Bruniales

[Bruniaceae + Columelliaceae]

Apiales

Pennantiaceae [Torricelliaceae [Griseliniaceae [Pittosporaceae [Araliaceae

[Myodocarpaceae + Apiaceae]]]]]

Paracryphiales

Paracryphiaceae

Dipsacales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

39

Adoxaceae [Diervillaceae [Caprifoliaceae [Linnaeaceae [Morinaceae [Dipsacaceae +

Valerianaceae]]]]]

LINKS TO ORDERS AND FAMILIES

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MAIN GROUPINGS

Asterids commelinids core eudicots asterid 1 asterid

2 eudicots gymnosperms Magnoliophyta monocots N-fixing claderosids seed plants

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Abietales Acanthales Acerales Acorales Actinidiales Actinostrobales Adoxales Aescul

ales Agavales Aizoales AkanialesAlismatales Alliales Alseuosmiales Alstroemeriales

Altingiales Amaranthales Amaryllidales Amborellales AmbrosialesAmmiales Amomal

es Ancistrocladales Anisophylleales Annonales Anthobolales Apiales Apocynales Apo

nogetonalesAquifoliales Arales Araliales Aralidiales Araucariales Arecales Aristolochi

ales Athrotaxidales Asarales AsclepiadalesAsparagales Asphodelales Asparagales Ast

eliales Atriplicales Aucubales Austrobaileyales Avenales

Balanitales Balanopales Balanophorales Balsaminales Barbeyales Barclayales Batales

Begoniales BerberidalesBerberidopsidales Betulales Biebersteiniales Bignoniales Bix

ales Boraginales Brassicales Brexiales Bromeliales BrunialesBrunoniales Burmannial

es Burserales Butomales Buxales Byblidales

Cactales Callitrichales Calycanthales Calycerales Campanulales Campynematales Can

ellales Cannales CapparalesCaprifoliales Cardiopteridales Carduales Caricales Carle

manniales Caryophyllales Cassiales Casuarinales CelastralesCentrolepidales Cephalot

ales Cephalotaxales Ceratophyllales Cercidiphyllales Chenopodiales Chironiales Chlor

anthalesChrysobalanales Cinchonales Circaeasterales Cistales Citrales Cocosales Col

chicales Columelliales CombretalesCommelinales Connarales Convolvulales Coriarial

es Cornales Corylales Corynocarpales Crassulales CrossosomatalesCucurbitales Cunn

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

40

inghamiales Cunoniales Cupressales Cycadales Cyclanthales Cymodoceales Cynarales

CynomorialesCyperales Cyrillales Cytinales

Daphnales Daphniphyllales Dasypogonales Datiscales Degeneriales Desfontainiales D

iapensiales Didymelales DillenialesDioales Dioncophyllales Dioscoreales Diospyrales

Dipentodontales Dipsacales Droserales

Ebenales Echiales Elaeagnales Elaeocarpales Elatinales Elodeales Emmotales Empetr

ales Ephedrales Ericales EriocaulalesErythropalaless Escalloniales Eucommiales Euph

orbiales Eupomatiales Eupteleales Euryalaless

Fabales Fagales Falcatifoliales Ficales Flacourtiales Flagellariales Fouquieriales Franc

oales Frangulales

Galiales Garryales Geissolomatales Gentianales Geraniales Gesneriales Ginkgoales G

laucidiales Globulariales GnetalesGoodeniales Greyiales Griseliniales Grossulariales

Grubbiales Gunnerales Gyrocarpales Gyrostemonales

Haemodorales Haloragales Hamamelidales Hanguanales Heisteriales Helleborales He

lwingiales Himantandrales HippuridalesHomaliales Hortensiales Huales Huerteales

Hydatellales Hydnorales Hydrangeales Hydrastidales HydrocharitalesHydropeltidales

Hydrostachyales Hypericales Hypoxidales

Icacinales Illiciales Iridales Irvingales Iteales Ixiales

Jasminales Juglandales Julianiales Juncaginales Juncales

Lacistematales Lactoridales Lamiales Lardizabalales Laurales Lecythidales Ledocarpal

es Leitneriales LentibularialesLigustrales Liliales Limnanthales Linales Liriales Loasal

es Lobeliales Loganiales Lonicerales Loranthales LowialesLythrales

Magnoliales Malpighiales Malvales Marathrales Marcgraviales Mayacales Medusagy

nales Medusandrales MelanthialesMelastomatales Meliales Melianthales Meliosmal

es Menispermales Menyanthales Metteniusales MitrastemonalesMiyoshiales Moni

miales Moringales Musales Myricales Myristicales Myrothamnales Myrsinales Myrta

les

Najadales Nandinales Narcissales Nartheciales Nelumbonales Nepenthales Neuradal

es Nitrariales Nolanales NothofagalesNyctaginales Nymphaeales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Ochnales Oenotherales Olacales Oleales Onagrales Oncothecales Opuntiales Orchid

ales Oxalidales

Saxifragales Pandales Pandanales Papaverales Paracryphiales Parasitaxales Paridales

Parnassiales PassifloralesPenaeales Pennantiales Pentaphragmatales Peridiscales P

etiveriales Petrosaviales Phellinales Philydrales PhyllanthalesPhysenales Phytolaccal

es Picramniales Pinales Pinguiculales Piperales Pittosporales Plantaginales Platanale

sPlumbaginales Poales Podocarpales Podophyllales Podostemales Polemoniales Pol

ygalales Polygonales PontederialesPortulacales Posidoniales Potamogetonales Primu

lales Proteales

Quercales Quillajales Quintiniales

Rafflesiales Ranunculales Rapateales Resedales Restionales Rhabdodendrales Rhamn

ales Rhinanthales RhizophoralesRhodorales Rhoipteleales Roridulales Rosales Rouss

eales Rubiales Ruppiales Rutales

Sabiales Salicales Salvadorales Samolales Samydales Sanguisorbales Santalales Sapin

dales Sapotales SarracenialesSaxegotheales Saxifragales Scheuchzeriales Sciadopitya

les Scleranthales Scrophulariales Scyphostegiales SedalesSilenales Simmondsiales S

milacales Solanales Sphenocleales Sphenostemonales Stangeriales Stellariales Stemo

nalesStilbales Stylidiales Styracales Surianales

Taccales Tamales Tamaricales Taxales Taxodiales Tecophilaeales Terebinthales Tern

stroemiales Theales TheligonalesThymelaeales Tiliales Tofieldiales Torricelliales Tov

ariales Tribelales Trilliales Trimeniales Triuridales TrochodendralesTropaeolales Turn

erales Typhales

Ulmales Urticales

Vacciniales Vahliales Vallisneriales Velloziales Veratrales Verbenales Viburnales Vinc

ales Violales Viscales VitalesVochysiales

Welwitschiales Winterales

Xanthorrhoeales Ximeniales Xyridales

Zamiales Zingiberales Zosterales Zygophyllales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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In some cases clicking on a family synonym will send you directly to the subfamily in

which the genus of the basionym is to be found

Abietaceae Abolbodaceae Abrophyllaceae Acalyphaceae Acanthaceae [Acanthochla

mydaceae] Acarnaceaee AceraceaeAchariaceae Achatocarpaceae Achradaceae Achy

ranthaceae Acmopylaceae Acoraceae Acristaceae ActaeaceaeActinidiaceae Actinota

ceae Amentotaxaceae Adenogrammaceae Adoxaceae Aegialitidaceae Aegicerataceae

Aegilopaceae Aeginetiaceaee Aegiphilaceae Aesculaceae Aextoxicaceae Agapanthac

eae Agavaceae AgdestidaceaeAgialidaceae Agrimoniaceae Agrostidaceae Ailanthace

ae Aitoniaceae Aizoaceae Akaniaceae Alangiaceae AlchemillaceaeAldrovandaceae A

lismataceae Alliaceae Allioniaceae Allophylaceae Aloaceae Alopecuraceae Alpiniacea

e AlseuosmiaceaeAlsinaceae Alsodeiaceae Alstroemeriaceae Altingiaceae Alzateacea

e Amaranthaceae Amaryllidaceae Amborellaceae Ambrosiaceae Amentotaxaceae A

mmanniaceae Ammiaceae Amomaceae Ampelidopsaceae Amygdalaceae Amyridacea

eAnacardiaceae Anagallidaceae Anarthriaceae Anchusaceae Ancistrocladaceae Andr

omedaceae AndropogonaceaeAndrostachyaceae Androsynaceae Anemarrhenaceae

Anemonaceae Angelicaceae Anisophylleaceae AnnonaceaeAnomochloaceae Anopter

aceae Anrederaceae Anthemidaceae Anthericaceae Anthobolaceae Antidesmataceae

Antirrhinaceae Antoniaceae Aparinaceae Aphanopetalaceae Aphloiaceae Aphyllanth

aceae Apiaceae ApocynaceaeApodanthaceae Aponogetonaceae Aporusaceae Apose

ridaceae Apostasiaceae Aptandraceae AquifoliaceaeAquilariaceae Aquilegiaceae A

raceae Aragoaceae Araliaceae Aralidiaceaee Araucariaceae Arbutaceae Arceuthidace

aeArceuthobiaceae Arctostaphylaceae Arctotidaceae Ardisiaceae Arecaceae Argoph

yllaceae Arisaraceae AristolochiaceaeAristoteliaceae Arjonaceae Armeriaceae Artem

isiaceae Arthrotaxidaceae Artocarpaceae Arundinaceae ArundinellaceaeAsaraceae A

sclepiadaceae Ascyraceaee Asiraceae Asparagaceae Asperulaceae Asphodelaceae As

pidistraceaeeAsteliaceae Asteraceae Asteranthaceae Asteropeiaceae Astragalaceae

Astrocarpaceae AthanasiaceaeAtherospermataceae Atriplicaceae Atropaceae Aucuba

ceae Aurantiaceae Austrobaileyaceae Austrotaxaceae AvenaceaeAverrhoaceae Avet

raceae Avicenniaceae Azaleaceae Azimaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Balanitaceae Balanopaceae Balanophoraceae Balsameaceae Balsaminaceae Bambusa

ceae Barbaceniaceae BarbeuiaceaeBarbeyaceae Barclayaceae Barringtoniaceae Base

llaceae Bataceae Baueraceae Bauhiniaceae Baxteriaceae BegoniaceaeBehniaceae B

elangeraceae Belloniaceae Bembiciaceae Berberidaceae Berberidopsidaceae Berryac

eaee BersamaceaeBertyaceae Berzeliaceae Besleriaceae Betaceae Betulaceae Biebe

rsteiniaceae Bifariaceae Bignoniaceae BischofiaceaeBixaceae Blakeaceae Blandfordia

ceae Blattiaceae Blepharocaryaceae Blitaceae Blyxaceae Boerlagellaceae Bolivaracea

eBombacaceae Bonnetiaceae Bontiaceae Boopidaceae Boraginaceae Borassaceae B

oroniaceae BoryaceaeBotryodendraceae Bougainvilleaceae Boweniaceae Brassicacea

e Brachycaulaceae Bretschneideraceae BrexiaceaeBromeliaceae Brownlowiaceae Br

unelliaceae Bruniaceae Brunoniaceae Brunsvigiaceae Bryoniaceae BuchneraceaeBuci

daceae Buddlejaceae Buglossaceae Bulbocodiaceae Bumeliaceae Bupleuraceae Burc

hardiaceae BurmanniaceaeBurseraceae Butneriaceae Butomaceae Buxaceae Byblida

ceae Byttneriaceae

Cabombaceae Cacaoaceae Cactaceae Caesalpiniaceae Caladiaceae Calamaceae Calce

olariaceae CalectasiaceaeCalendulaceae Callaceae Callicomaceae Calligonaceae Calli

traceae Callitrichaceae Calochortaceae CalophyllaceaeCalthaceae Calycanthaceae Ca

lyceraceae Cambogiaceae Camelliaceae Campanulaceae CampynemataceaeCanacom

yricaceae Canellaceae Cannabaceae Cannaceae Canopodaceae Canotiaceae Cansjera

ceae CapparaceaeCaprariaceae Caprifoliaceae Cardiopteridaceae Carduaceae Carica

ceae Carissaceae Carlemanniaceae CarpinaceaeCarpodetaceae Cartonemataceae Ca

ryocaraceae Caryophyllaceae Casearieaceae Cassiaceae CassipoureaceaeCassythacea

e Castaneaceae Castelaceae Casuarinaceae Catesbaeaceae Cathedraceae Cecropiace

ae CedraceaeCedrelaceae Celastraceae Celosiaceae Celtidaceae Centaureaceae Cen

trolepidaceae Centroplacaceae CepaceaeCephalanthaceae Cephalotaceae Cephalota

xaceae Cerastiaceae Ceratoniaceae Ceratophyllaceae CerberaceaeCercidiphyllaceae

Cercocarpaceae Cercodiaceae Cereaceae Cerinthaceae Ceroxylaceae Cestraceae Cev

alliaceaeChailletiaceae Chamaedoraceae Chamaemoraceae Chamelauciaceae Chaun

ochitonaceae Chelidoniaceae ChelonaceaeChenopodiaceae Chimonanthaceae Chingi

thamnaceae Chionographidaceae Chiranthodendraceae ChironiaceaeChlaenaceae Ch

loanthaceae Chloranthaceae Chloridaceae Chlorogalaceae Chrysobalanaceae Cicerac

eae CichoriaceaeCimicifugaceae Cinchoniaceae Circaeaceae Circaeasteraceae Cissac

eae Cistaceae Citraceae Clematidaceae CleomaceaeClethraceae Clusiaceae Cneorac

eae Cnestidaceae Cnicaceae Cobaeaceae Cochlospermaceae Cocosaceae Coffeaceae

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Colchicaceae Coleogynaceae Columelliaceae Combretaceae Commelinaceae Comocl

adiaceae Compositae CompsoaceaeConantheraceae Connaraceae Conostylidaceae C

onvallariaceae Convolvulaceae Cordiaceae CoreopsidaceaeCoriandraceae Coriariacea

e Coridaceae Corispermaceae Cornaceae Corokiaceae Coronillaceae Correaceae Cor

rigiolaceaeCorsiaceae Corydalaceae Corylaceae Corynocarpaceae Coryphaceae Cost

aceae Cotyledonaceae CoulaceaeCoutariaceae Coutoubiaceae Crassulaceae Crescen

tiaceae Cressaceae Crinaceae Crocaceae CroomiaceaeCrossosomataceae Crotonacea

e Cruciferae Cryptaceae Crypteroniaceae Cryptocorynaceae CryptomeriaceaeCtenol

ophonaceae Cucurbitaceae Cunninghamiaceae Cunoniaceae Cupressaceae Curcumac

eae Curtisiaceae CuscutaceaeCyananthaceae Cyanastraceae Cynanchaceae Cyanella

ceae Cycadaceae Cyclanthaceae CyclantheraceaeCyclocheilaceae Cydoniaceae Cymo

doceaceae Cynaraceae Cynocrambaceae Cynomoriaceae Cyperaceae CyphiaceaeCyp

hocarpaceae Cypripediaceae Cyrillaceae Cyrtandraceae Cyrtanthaceae Cytinaceae

Dacrycarpaceae Dactylanthaceae Damasoniaceae Daphnaceae Daphniphyllaceae Das

ypogonaceae DatiscaceaeDaturaceae Daucaceae Davidiaceae Davidsoniaceae Decais

neaceae Deeringiaceae Degeneriaceae DelphiniaceaeDendrophthoaceae Columelliac

eae Detariaceae Dialypetalanthaceae Dianellaceae Dianthaceae DiapensiaceaeDicha

petalaceae Dichondraceae Diclidantheraceae Dicrastylidiaceae Dictamnaceae Didiere

aceae DidymelaceaeDidymocarpaceae Diegodendraceae Diervillaceae Digitalidaceae

e Dilatridaceae Dilleniaceae Dioaceae DionaeaceaeDioncophyllaceae Dioscoreaceae

Diosmaceae Diospyraceae Dipentodontaceae Diphylleiaceae Diplolaenaceae Dipsaca

ceaeDipterocarpaceae Dirachmaceae Disanthaceae Diselmaceae Dodonaeaceae Do

mbeyaceae Donatiaceae DorsteniaceaeDortmannaceae Doryanthaceae Drabaceae D

racaenaceae Dracontiaceae Drimyidaceae Droseraceae DrosophyllaceaeDryadaceae

Duabangaceae Duckeodendraceae Dulongiaceae Durantaceae Durionaceae Dysphani

aceae

Ebenaceae Eccremidaceae Ecdeiocoleaceae Echiaceae Echinopaceae Ehretiaceae Ela

eagnaceae ElaeocarpaceaeElegiaceae Elatinaceae Ellisiaceae Ellisiophyllaceae Elodea

ceae Elytranthaceae Embeliaceae Emblingiaceae EmmotaceaeEmpetraceae Encepha

lartaceae Engelhardtiaceae Enhalaceae Epacridaceae Ephedraceae Ephemeranthacea

eEpilobiaceae Epimediaceae Eragrostidaceae Eremolepidaceae Eremosynaceae Erica

ceae Erinaceae EriocaulaceaeEriogonaceae Eriospermaceae Erodiaceae Erycibaceae

Eryngiaceae Erysimaceae Erythroniaceae ErythropalaceaeErythrospermaceae Erythro

xylaceae Escalloniaceae Eschscholziaceae Eucomidaceae Eucommiaceae Eucryphiace

aeEuonymaceae Eupatoriaceae Euphorbiaceae Euphrasiaceae Euphroniaceae Eupo

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matiaceae Eupteleaceae EuryalaceaeEustrephaceae Euthemidaceae Eutocaceae Exb

ucklandiaceae Exocarpaceae

Fabaceae Fagaceae Falcatifoliaceae Ferulaceae Festucaceae Ficaceae Ficoidaceae Fit

zroyaceae FlacourtiaceaeFlagellariaceae Flindersiaceae Foetidiaceae Forestieraceae

Fothergillaceae Fouquieriaceae Fragariaceae FrancoaceaeFrangulaceae Frankeniacea

e Fraxinaceae Fraxinellaceae Freycinetiaceae Fritillariaceae Fuchsiaceae Fumariacea

eFunkiaceae

Gaiadendraceae Galacaceae Galanthaceae Galaxiaceae Galedupaceae Galeniaceae G

aliaceae Garciniaceae GardeniaceaeGardneriaceae Garryaceae Geissolomataceae Ge

itonoplesiaceae Gelsemiaceae Geniostomaceae GentianaceaeGeonomataceae Geosir

idaceae Geraniaceae Gerrardinaceae Gesneriaceae Gethyllidaceae Gilliesiaceae Gina

lloaceaeGinkgoaceae Gisekiaceae Gladiolaceae Glaucidiaceae Glechomaceae Glinace

ae Globulariaceae Gnaphaliaceae GnetaceaeGoetzeaceae Gomortegaceae Gomphiac

eae Gomphrenaceae Gonystylaceae Goodeniaceae Gordoniaceae GouaniaceaeGoup

iaceae Gramineae Gratiolaceae Grewiaceae Greyiaceae Grielaceae Griseliniaceae Gr

onoviaceae GrossulariaceaeGrubbiaceae Guaiacanaceae Guamatelaceae Guettardac

eae Gunneraceae Gustaviaceae Guttiferae GyrocarpaceaeGyrostemonaceae

Hachetteaceae Haemanthaceae Haemodoraceae Halesiaceae Hallieraceae Halocarpa

ceae Halophilaceae HalophytaceaeHaloragaceae Hamamelidaceae Hanguanaceae H

aptanthaceae Harmandaceae Hebenstretiaceae HectorellaceaeHederaceae Hedyosm

aceae Hedyotidaceae Hedysaraceae Heisteriaceae Heleniaceae Heliamphoraceae H

elianthaceaeHelianthemaceae Helichrysaceae Heliconiaceae Helicteraceae Heliotropi

aceae Helleboraceae Heloniadaceae HelosidaceaeHelwingiaceae Hemerocallidaceae

Hemimeridaceae Hemiphylacaceae Henriqueziaceae Henslowiaceae Hermanniaceae

Hernandiaceae Herniariaceae Herreriaceae Hesperocallidaceae Heterantheraceae He

teropyxidaceae HeterostylaceaeHibbertiaceae Hibiscaceae Hilleriaceae Himantandra

ceae Hippocastanaceae Hippocrateaceae HippomanaceaeHippophaeaceae Hippurida

ceae Hirtellaceae Holacanthaceae Homaliaceae Hopkinsiaceae Hoplestigmataceae H

ordeaceaeHornschuchiaceae Hortensiaceae Hortoniaceae Hostaceae Houstoniaceae

Huaceae Huerteaceae HugoniaceaeHumbertiaceae Humiriaceae Hyacinthaceae Hyd

atellaceae Hydnoraceae Hydrangeaceae Hydrastidaceae HydrillaceaeHydrocharitacea

e Hydrocotylaceae Hydrogetonaceae Hydroleaceae Hydropeltidaceae Hydrophylacac

eae HydrophyllaceaeHydrostachyaceae Hymenocardiaceae Hypecoaceae Hyperanth

eraceae Hypericaceae Hypopityaceae HypoxidaceaeHypseocharitaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta

ceae ImperatoriaceaeInocarpaceae Inulaceae Ionidiaceae Iriarteaceae Iridaceae Irvi

ngiaceae Isnardiaceae Isophysidaceae IteaceaeStrasburgeriaceae Ixiaceae Ixioliriace

ae Ixonanthaceae

Jalapaceae Jamboliferaceae Japonoliriaceae Jasionaceae Jasminiaceae Johnsoniaceae

Joinvilleaceae JuglandaceaeJulianiaceae Juncaceae Juncaginaceae Juniperaceae Jus

siaeaceae Justiciaceae

Kadsuraceae Kaliphoraceae Kalmiaceae Kaniaceae Kiggelariaceae Kingdoniaceae Kin

giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae

Krameriaceae

Labiatae Lacandoniaceae Lachenaliaceae Lacistemataceae Lactoridaceae Lactucaceae

Lagerstroemiaceaee LamiaceaeLampsanaceae Lanariaceae Langsdorffiaceae Lantan

aceae Lapageriaceae Lardizabalaceae Lasiopetalaceae LauraceaeLawsoniaceae Laxm

anniaceae Lecythidaceae Ledaceae Ledocarpaceae Leeaceae Leguminosae Leitneriac

eae LemnaceaeLennoaceae Lentibulariaceae Lentiscaceae Leoniaceae Leonticaceae

Lepidobotryaceae LepidocarpaceaeLepidocaryaceae Lepidocerataceae Leptaulaceae

Leptospermaceae Lepturaceae Lepuropetalaceae LeuctenbergiaceaeLibocedraceae

Licaniaceae Ligustraceae Lilaeaceae Liliaceae Limeaceae Limnanthaceae Limnocharit

aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace

ae Linderniaceae Lindleyaceae Linnaeaceae LippayaceaeLiriaceae Liriodendraceae Li

ssocarpaceae Littorellaceae Loasaceae Lobeliaceaee Loganiaceae LomandraceaeLoni

ceraceae Lopeziaceae Lophiocarpaceae Lophiolaceae Lophiraceae Lophophytaceae L

ophopyxidaceae LoranthaceaeLotaceae Lowiaceae Lupulaceaee Luxemburgiaceae Lu

zuriagaceae Lychnidaceae Lyciaceae LyginiaceaeLygodisodeaceae Lysimachiaceae Ly

thraceae

Macarisiaceae Mackinlayaceae Madiaceae Maesaceae Magnoliaceae Malaceae Male

sherbiaceae MalortieaceaeMalpighiaceae Malvaceae Mangiaceae Manicariaceae Ma

paniaceae Marantaceae Marathraceae MarcgraviaceaeMartyniaceae Mastixiaceae

Matricariaceae Maundiaceae Mayacaceae Medeolaceae Medusagynaceae Peridiscac

eaeMelaleucaceae Melampyraceae Melanophyllaceae Melanthiaceae Melastomatac

eae Meliaceae Melianthaceae MelicaceaeMeliosmaceae Melittidaceae Melochiacea

e Memecylaceae Mendonciaceae Menispermaceae Menthaceae MenyanthaceaeMe

nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

Nypaceae Nyssaceae

Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

ceae Papilionaceae Pappophoraceae PapyraceaeParacryphiaceae Parasitaxaceae Par

ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

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hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

49

cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

50

ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 8: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

8

particular is very well established In a somewhat similar situation if Lemna and its

relatives are a clade sister to most other Araceae should they be recognised as a

separate family Gymnostachys a phenetically fairly distinctive taxon as well as the less

phenetically distinct Orontioideae would have to be recognised as a separate families

(or combined as a single family) too but Araceae in a somewhat restricted sense would

be somewhat more morphologically coherent although not greatly so and not notably

distinct However it is in the very nature of such decisions to be somewhat arbitrary

and unsatisfactory hence the emphasis on consensus classifications here and on

classifications as simply being a means to an end

A useful distinction can be drawn between crown groups and stem groups The

former are monophyletic and include the extant members of a clade and their

immediate common ancestor (see the figure below) The groups characterized in this

site are such groups Thus Proteaceae here are crown group Proteaceae apomorphies

like the single carpel four-merous perianth etc being found in this common ancestor

Stem groups on the other hand include all the members of a lineage immediately after

its split from its sister group and all branches of this lineage In the case of Proteaceae

the stem group would include everything after its split from its sister group

Platanaceae Obviously most of the organisms in the lineage that terminates in crown-

group Proteaceae are unknown only a few being known as fossils and it is not known

exactly where particular apomorphies of crown group Proteaceae evolved along this

lineage

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

9

Indeed there is no reason other than convention or convenience why any group

should not be segregated into several smaller monophyletic groups or merged to

produce a larger unit we can talk about one large thing or about several smaller things

Thinking about aspects of size findings in ethnobiology and cognitive psychology can be

used to suggest that a moderate number - probably fewer than 500 - of families is a

reasonable goal at which to aim and that groupings of taxa throughout any system

should be rather small in size (eg Berlin 1992 Stevens 1994 1997) Major systems such

as those of Linnaeus and Bentham and Hooker were constructed explicitly so as to ease

the burden on the memory (Stevens 1997 2002 see also Scharf 2007) the latter in

particular ensuring that all groups in their classification were relatively small often

containing three to eight immediately subordinate taxa - but by no means all their

groups were formally named Along the same lines Burtt (1977b) suggested that the

number of names at any rank should be at most one third those at the immediately

lower rank - and monotypic taxa might not need a formal name Consistent with such

ideas a fairly broad view of families and orders is taken here whenever the constraints

of monophyly and other criteria used when constructing classifications (see above)

permit The APG system is thus a convention consistent with ideas such as those

expressed by Backlund and Bremer (1998) for similar conventions see eg the Grass

Phylogeny Working Group (2001) and for fungi Hibbett et al (2007)

There are other views Thus Takhtajan (1997) has suggested that smaller families

are more natural This is incorrect Monophyletic groups that include fewer taxa -

Takhtajans smaller families - do not necessarily have more apomorphies than larger

groups even if members of smaller groups will always have more features in general in

common than the larger group that includes them That is they will have their

apomorphies their unique features as well as progressively more plesiomorphies

features found both in the small groups and in the larger clades of which they are a part

having more features in common is indeed one common meaning of more natural

(Note that the implication of the word natural has long been a group of the kind

[usually unspecified] that I think should be recognised and if other botanists had

different ideas too bad their groups were necessarily not natural [see Bather 1927] It

is thus rarely a helpful word and is not used here) Furthermore if this approach is

adopted we will find a slippery slope ahead By this kind of argument all families should

be very small since their members will have a great deal in common and so will be

most natural However as families (for example) are split the relationships that are

evident between the segregates and that were responsible for their being placed in a

single family in the first place will seem to necessitate the recognition of a new order

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

10

etc as is evident in Takhtajans own work - general taxonomic inflation is the result (see

also comparable suggestions in a cladistic context for Brassicales in particular - Ronse de

Craene amp Haston 2006) Such splitting is also questionable when teaching and learning

families since the student needs to understand the system as a whole However for

some genera removed from the families that until now have included them the

phenetic-classificatory-phylogenetic structure in their new home may mandate the

recognition of small families On the other hand Takhtajans suggestion that narrowly

defined families are more useful for phylogenetic studies may be true Indeed I have

more than once regretted prematurely combining groups whether species (in the

context of monographic work) or families (in the course of preparing these notes) but

this is largely a separate issue

Van Steenis (1978) Philipson (1987b) and others have rightly questioned the value

of splitting a group when ideas of the relationships of its constituent members have not

changed - that is very good reasons have to be provided for splitting a family if the

genera within it remain part of the same clade rather than belonging to another clade

Thus APG (2003) broadened the circumscription of Malvaceae because of the

parapolyphyly of some of the families that had historically been associated with it (Judd

amp Manchester 1997 Alverson et al 1999 Bayer et al 1999) These families particularly

Tiliaceae and Sterculiaceae were not at all easy to distinguish their close relationship

(see eg Brown 1814) and overall similarity had long been conceded and to some

workers at least their combination has come as something of a relief Although most of

the larger clades within Malvaceae sl remain difficult to distinguish even with flowers

Cheek (2007) opts for a wholesale and novel dismemberment into ten families

however the very good reasons for doing this are wanting

The same principles are of course applicable when it comes to dividing genera little

other than a headache is gained by splitting genera such

as Drosera and Gnetum (Doweld 2000) as has recently been proposed Thus if an

established genus divides into two (or more) clades this is not a signal for recognising

two groups at the same level - so here it could be argued that the dismemberment

of Pterostylis (Jones amp Clements 2002b) was somewhat unfortunate Along the same

lines if a newly-discovered taxon is sister to an existing named taxon - say a genus - this

does not necessitate the description of a separate genus for the newly described species

(cf Davis 2002) (Of course some [eg Thorne 1976] have suggested that the sizes of

gaps between groups at the same rank should be similar but any principle like this is

inherently flawed since morphological gaps are more unstable than phylogenetic

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

11

relationships and applying it across all flowering plants would both be difficult and cause

substantial changes - see also below) Humphreys and Linder (2009) provide a well-

documented survey of generic concepts in plants which the reader should consult they

note that generic limits (broad versus narrow) have oscillated historically and that

currently larger genera tend to be recognised because studies tend to be on a broader

scale than in the past

Note that invoking similarity or difference - whether qualified (considerable

similarities substantial differences) or not - in a cladistic context as justification for

combining or splitting taxa is not a particularly strong argument (see eg

CardiopteridaceaeStemonuraceae - Karingrehed 2002c) Similarity and difference can

neither be defined precisely since what may seem to be substantial similarities to me

may not to the next person nor are they likely to be stable in the face of our changing

knowledge of morphology and what might be synapomorphies

I might have prefered to merge some families recognised here or split others but

by and large I do not think my own preferences matter very much - and I take the same

position with regards to comparable preferences expressed by others Indeed the

bottom line is that in flagged hierarchies of the kind used here the limits of any

monophyletic unit generally taught and discussed particularly other than species can

be established only by convention and consensus (eg Stevens 2002 2006a Entwisle amp

Weston 2005) This is rather different from the reasonable observation that the

phylogeny itself cannot be achieved by consensus (Thorne 1976) for in most cases there

is a fact of the matter when it comes to relationships Given the increasing support for

the outlines of angiosperm phylogeny a stable consensus classification based on this

phylogeny seems attainable Indeed in addition to providing current ideas of

relationships of seed plants in a synthesised form this site is part of an attempt to build

such a consensus about the circumscription of taxa (see APG 1999 2003 2009 Grass

Phylogeny Working Group 2001 Mabberley 2008) Reaching such a consensus is vital

since what we know of angiosperm phylogeny allows a very large number of

classifications to be based on it and as we find out more the number of possible

classifications increases greatly Unfortunately however nature does not dictate what

the classification should be All classifications are constructed by humans to

communicate particular aspects of groups and relationships Our goals as systematists

are surely to produce robust hypotheses of relationships to understand the evolution of

morphology and the like - but not to argue ad nauseam whether something should be

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

12

a family or a subfamily That way surely lies madness and worse the discredit of our

discipline

There are similar issues whatever naming system is used Thus in phylogenetic

naming (Baum et al 1998 for an example but cf Baum et al 2004 for the PhyloCode

see Cantino amp De Queiroz 2006) an unflagged hierarchy is used in which any

terminations of names used are uninformative about the relative position of taxa If one

adopts the principle of phylogenetic naming one indeed does not have to worry about

the nomenclatural consequences caused by lumping or splitting any well-supported

clade can be named without affecting the name of more or less inclusive clades

Unfortunately unflagged hierarchies have very serious deficiences as communication

devices because they lack one aspect essential in language biological or otherwise -

they contain no intrinsic information about the relationships of the group in question to

others (eg Pfeil amp Crisp 2005 Stevens 2006a) Recent suggestions for using prefixes like

Apo- and Pan- to PhyloCode names will however allow limited information of this

kind to be conveyed but only as it pertains to individual branches and current

proposals do not even mandate that the prefixes be employed consistently In any

event such proposals simply prevent the potential tripling of the number of quite

different names used to describe different aspects of a phylogenetic tree over those

used to name monophyletic groups pure and simple In general where n is the number

of extant species in a group the number of clades in such a group = n-1 (Species will

also need names too for their names see Dayrat et al 2008) Importantly here too

consensus over the clade names commonly learned by students and used in herbaria is

needed otherwise communication will be impeded the names themselves will provide

no guidelines as to which should be chosen The situation is of course more complicated

than this Terminations that convey ideas of rank in a phylogenetic classification can also

be used in phylocode names - however there they will carry no implications of rank

How they will be used is another matter of course

Of course there are other philosophies of classification and some still prefer

evolutionary classifications There classificatory principles differ substantially from those

followed here eg the recognition of paraphyletic taxa may be permitted however

detailed reasons for prefering the taxa that are recognised are rarely given although

nature and natural groups are often mentioned (cf Stuessy amp Koumlnig 2008) For

summaries of commonly used systems see Brummitt (1992) and Mabberley (2008)

new evolutionary systems appear every year or so However even those who allow or

promote the recognition of paraphyletic groups (eg Grant 2003 Thorne 2007

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

13

Heywood et al 2007) may find it of interest to examine a system recognizing only

monophyletic groups it provides a rather different understanding of evolution

Evolutionary classifications in general try and combine phylogeny and

morphological gaps although that is no easy thing to do - it is akin to combining chalk

and cheese (for an attempt to make this impossible task seem to be more objective see

Stuessy amp Koumlnig 2008) However there is a principle from evolutionary classification

that is relevant and I think quite useful here The size of the gap between two groups

tends to be inversely proportional to the sizes of the groups involved (Davis amp Heywood

1963) One can imagine a situation in which a large group is formally divided even

although the distinguishing characters of the two are weak whereas a smaller group

similarly divisable will be left intact

To summarize If hypotheses of phylogeny remain stable we should be able to base

a stable classification on that phylogeny and then get on with our work that is testing

the phylogenies we have elucidating phylogenies in areas where relationships are

unclear studying the evolution of morphology describing species etc In this context

the spread of the Angiosperm Phylogeny Group system (see below) and its widespread

utilisation in technical literature also floras (eg van der Meijden 2005) textbooks (eg

Simpson 2006 Judd et al 2007 [third edition]) dictionaries (Mabberley 2008) more

popular literature (eg Souza amp Lorenzi 2005 Spears 2006) and as an outline for a new

herbarium sequence (Haston et al 2007) is gratifying The posibility that one might be

able to develop a stable phylogeny-based classification of families and in particular

orders represents a dramatic turn-around from the pessimistic attitude about such

higher-level groupings expressed by Davis and Heywood (1963) and Thorne (1976) the

latter even suggesting that we should bury forever the metaphor of the phylogenetic

tree as highly unrealistic (ibid p 56) Returning to Godfray and Knapps (2004) users

of classifications who want a stable informative and accessible classification that

enables easy identification - unfortunately they want cake with everything and cannot

get it - these pages attempt to satisfy as many of their needs as possible but phylogeny

and monophyly are the primary shapers of the APG classification

On this classification in particular

Here I very largely follow the Angiosperm Phylogeny Group classification (APG

2003) Any differences are not to be interpreted as differences in principle simply that

new phylogenies continue to be published and here I attempt to provide an overview

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

14

of current ideas of higher-level relationships of seed plants The Angiosperm Phylogeny

Group classification is based on well-supported relationships evident in the numerous

molecular studies that began to appear in the late 1980s (see APG 1999 for the

principles underlying the classification) but recent phylogenetic work does not

contradict the major outlines of the trees used by APG II (2003) or even those in APG

I (APG 1999) The APG classification has for the most part been conservative and I

am similarly conservative here For the names and authorities I follow APG III (2009)

although the names of the authors may not always be correct these being in something

of a state of flux for the superordinal names used see Chase and Reveal (2009)

although classification at this level here is not exhaustive - only clades with several

orders have superordinal names

Some changes in our ideas of relationships and hence in the clades we talk about

are particularly likely in parts of Lamiales Caryophyllales and Malpighiales for example

although this will not affect the orders themselves Thus some changes to clade

circumscriptions are to be expected although I do not expect them to be substantial

but changes are neither a defect of cladistics nor a necessary consequence of the use of

molecular data Clades are hypotheses of relationships and as hypotheses they may be

overturned However work since APG I has not suggested other than minor changes

in the compositions of the orders even if the odd genus or even family is turning out to

be seriously misplaced - recent examples are Hydatellaceae (from monocots-Poales to

Nymphaeales Saarela et al 2007) Guamatelaceae (from Rosales-Rosaceae to

Crossosomatales Oh amp Potter 2006) and Perrottetia and Bhesa (from Celastrales-

Celastraceae to Huerteales and Malpighiales respectively Zhang amp Simmons 2006) The

main changes have been clarification of the relationhips of individual families or groups

of families that were of uncertain position eg of Chloranthaceae (Moore et al 2007)

Ceratophyllaceae (Jansen et al 2007) and unplaced asterid II families (Winkworth et al

2008a) As mentioned above phylogenies in the area of main interest in these pages are

overwhelmingly tree-like Although there is evidence for acquisition of host plant genes

by parasites (see below) even wider but inexplicable transfer of mitochochondrial

genes in plants like Amborella (Bergthorsson et al 2004 cf Goremykin et al 2009a)

and even transfers of nuclear genes (Vallenback et al 2008) these are the exception

rather than the rule

In cases where the Angiosperm Phylogeny Group II allowed alternatives as to the

limits of families - Papaveraceae in the broad sense or Papaveraceae plus

Pteridophyllaceae plus Fumariaceae Proteaceae in the broad sense or Proteaceae plus

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

15

Platanaceae - the choices made here follow common usage eg as in textbooks like

Judd et al (2007) and Simpson (2006) and particularly in the new edition of

Mabberleys The Plant Book (Mabberley 2008) This latter is itself an attempt to reflect a

consensus the result of taking the opinions of botanists at several meetings A largely

similar consensus is reflected in the APG III classification (APG 2009) where you will

see that alternative classifications have been dispensed with For many people the

existence of such alternative classifications will simply confuse so agreement over

which groupings to use when alternatives are permitted and then dispensing with the

alternatives that are not commonly used seems reasonable I am making the few

changes that are needed to bring the classification here in line with these others but

only gradually since there is not enough time in each day

As already mentioned higher-level relationships in general and the composition of

orders in particular have always presented something of a challenge to systematists

(eg Davis amp Heywood 1963) That being said the composition of some clades like

Apiales Crossosomatales and Pandanales is decidedly unexpected However it is

interesting to see that these higher level clades are generally accepted even in works

with different classificatory philosophies (eg Heywood et al 2007) For clades like

Malpighiales many of the family groupings within Asparagales etc attempts to find

distinctive characters have largely failed (but see Endress amp Matthews 2006a also

the Apomorphies page here) Interestingly as with families some groupings suggested

by molecular studies are supported by morphological andor chemical characters that

have long been known sometimes for over a hundred years the relationship between

Pittosporaceae and ApiaceaeAraliaceae are a case in point (Hegnauer 1969b and

references) As our knowledge of morphology and chemistry improves we can hope for

improvements in the characterisations of clades at all levels

In many cases the new family limits of the Angiosperm Phylogeny Group (see

APG 1999 2003) are not really controversial although changes from the limits

commonly accepted only a decade ago are sometimes quite dramatic (eg Wagenitz

1997) Thus the split of the old Saxifragaceae sl is necessitated by its extreme

polyphyly as also with Icacinaceaesl and Cornaceae sl However the limits and

relationships of such groups had long been considered to be unsatisfactory but until

recently there had been no real reason to prefer one proposed arrangement over

another It is generally accepted that the limits of Lamiaceae and Verbenaceae have to

be redrawn although there is as yet no compelling evidence that the redrawn taxa are

not sister taxa (there is no evidence that they are) But whatever their relationships the

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

16

content of the clades has changed considerably and incidentally they are now easier to

identify than before the decision to recognise two families is not difficult The same is

true for Salicaceae and Achariaceae (Malpighiales) two previously small families that

have received the bulk of the old Flacourtiaceae Clade and hence taxon limits remain

difficult around Theaceae (Ericalessee alsoSladeniaceae Pentaphylacaceae)

and Euphorbiaceae (Malpighiales see

also Phyllanthaceae Picrodendraceae Putranjivaceae etc) Here current groupings

may not represent quite such dramatic changes in our understanding of relationships

Indeed parts of the old Euphorbiaceae that were separated may yet go back together

particularly Phyllanthaceae and Picrodendraceae (see Wurdack et al 2004 Davis et al

2005) even if Theaceae should indeed be dismembered (Geuten et al 2004) However

given that there is no molecular evidence that currently warrants combining all the

segregates of Euphorbiaceae and even if some do come together the clades the

families represent suggest novel groupings not recognised in current classifications

Furthermore Rafflesiaceae appear to be embedded within Euphorbiaceae s str (Davis

et al 2007) so the family is divided Of course maintaining even a moderately narrowly-

circumscribed Euphorbiaceae would entail reducing the iconic Rafflesiaceae to

synonymy Relationships in core Caryophyllales especially

around Phytolaccaceae andNyctaginaceae and also Portulacaceae are incompletely

understood and refashioning of taxon limits will doubtless be needed as cladistic

relationships become apparent (eg see Nyffeler 2007 Ogburn 2007a b) Some

groupings in the old Icacinaceaeand Olacaceae also remain unclear

The discovery of the relationships of parasitic and aquatic groups have presented a

particular challenge to systematists Morphologically some of these plants are so highly

modified that interpretation of the plant body in conventional terms is difficult or even

impossible Thus parasitic groups (broad-leaved angiosperms only) such as Rafflesiaceae

are hard to place since both the vegetative body and the flowers are changed almost

beyond recognition (flowers of rafflesiaceae are a good example of gigantism - Davis et

al 2007) Furthermore plastid gene sequences may be difficult or impossible to obtain

the chloroplast DNA in particular being highly degraded and the rate of molecular

change in general may be high so the problem of long-branch attraction is serious (eg

Duff amp Nickrent 1997 Nickrent et al 1998 Caddick et al 2002a G Petersen et al

2006b) Echlorophyllous sapromycoheterotrophic taxa mostly known from monocots

present similar problems However progress is being made placements for

Apodanthaceae Rafflesiaceae Mitrastemonaceae Cytinaceae and Cynomoriaceae have

recently been suggested (Barkman et al 2004 2007 Davis amp Wurdack 2004 Nickrent et

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

17

al 2004 Davis et al 2007) it seems likely that Burmanniaceae sl are polyphyletic but

both parts are in Dioscoreales (Merckx et al 2006) and relationships within the largely

hemiparasitic Santalales are also gradually being clarified (Maleacutecot 2002) The inclusion

of parasitic taxa in general molecular analyses can cause conniptions (eg Nickrent et al

2004 Davis et al 2004 Chase et al 2006 G Petersen et al 2006b) and there can be

horizontal transmission of genes (eg Davis amp Wurdack 2005 Vitaceae to Rafflesiaceae

Barkman et al 2007 the mitochondrial atp1 gene commonly moves)

Similarly in water plants neither vessels in particular nor much xylem in general is

needed leaves are highly modified and water-mediated pollination if adopted may

well be associated with major changes in floral morphology Here too recent molecular

studies suggest that aquatic groups with hitherto problematic relationships may find

homes Thus Podostemaceae are close to Clusiaceae and Calophyllaceae and are sister

to Hypericaceae (Malpighiales Kita amp Kato 2001) Hydatellaceae which used to be in

Poales are part of Nymphaeales (Saarela et al 2007) and Hydrostachyaceae may be

close to Hydrangeaceae (Cornales Xiang et al 2002) or perhaps should be placed in

Lamiales note that in the first two cases in particular there are morphological and

chemical features that support such a move If Podostemaceae are indeed close to

Clusiaceae I look forward to seeing hypotheses to explain how the dramatic changes in

the vegetative body that have made Podostemaceae so problematic for generations of

systematists took place That conventional wisdom has trouble in understanding or

explaining how the morphologies of groups like Clusiaceae and Podostemaceae can be

related is largely a problem with conventional wisdom Ceratophyllaceae seem to be

finding a position as sister to eudicots (see Moore et al 2007) but their morphology is

so derived that there is no morphological evidence of which I am aware for this

relationship There are also similar although less extreme morphological problems with

many plants pollinated by wind

In such situations especially with some water plants mycoheterotrophs and

parasites a variety of characters may be affected and particularly under such

circumstances groups have tended to be formed using a few or even only a single

character that seems to provide evidence of relationships Add to this the tendency to

weight some characters particularly strongly on a priori grounds and the result was

either the recognition of conglomerate taxa such as Amentiferae which now appear to

be highly polyphyletic or the segregation of families like Plantaginaceae s str (now

much expanded) and Leitneriaceae (now in Simaroubaceae) - all of these are more or

less wind pollinated

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

18

Even although relationships of many parasitic and aquatic groups are being

clarified and at least some of the proposed phylogenetic positions seem to be well

supported it is indeed often still very difficult to understand how such groups have

evolved from their more morphologically conventional relatives However we have had

all sorts of preconceptions as to how characters should evolve Thus we used to

assume that features like highly scalariform vessel perforation plates or the complete

absence of vessels or a flower with an androecium that had many stamens a superior

ovary or separate petals were necessarily plesiomorphic or primitive and conversely

a simple perforation plate and the presence of vessels an androecium with few

stamens inferior ovary or petals that were connnate were almost necessarily

apomorphic or advanced (but cf eg Stebbins 1951) Such assumptions are incorrect

(eg Soltis et al 2005b) Carpels may become secondarily free carpels may fail to close

the seeds then developing outside the confines of the carpel as in some Aspagaraceae-

Nolinoideae Violaceae Berberidaceae Malvaceae-Sterculioideae etc) in Peliosanthes

teta perhaps the only species in Peliosanthes(Asparagaceae-Nolinoideae) the ovary

varies from superior to inferior (Jessop 1976 species limits here need close

investigation see also Kuzoff et al 2001 and Soltis amp Hufford 2002 Saxifragaceae

Apiales etc) many-seeded carpels can evolve from few-seeded carpels

(Razafimandimbison et al 2008) monoecy may be derived from dioecy (Schaefer amp

Renner 2010 and references) Classic studies such as those by Babcock (eg 1947)

on Crepis that assumed that evolution - in this case of the karyotype in particular - was

unidirectional have needed comprehensive re-evaluation (Enke amp Gemeinholzer 2008)

Most if not all characters have reversed andor evolved in parallel as is clear in the

discussion of ovary position here especially in Poales Saxifragales and Asparagales and

also in comments on such morphologically distinctive taxa (when compared with their

immediate relatives) such as Menyanthaceae and especially Pittosporaceae Parallel

evolution may occur even at the level of amino acid substitution as in the independent

acquisition of the phosphoenolpyruvate carboxylase (pepC) gene in C4 photosynthesis in

grasses (Christin et al 2007b see also Blaumlsing et al 2000)

Thinking of how such characters have been used in the past in classifications at the

generic level the results have been similar to those just discussed above for parasitic

plants and others Relying too much on animal pollination syndromes to mark generic

boundaries has all too often led to taxa that are highly unsatisfactory phylogenetically

(see eg Acanthaceae Bignoniaceae Campanulaceae Ericaceae Lamiaceae

Orchidaceae) and over-reliance on characters of fruit and seed (see eg Brassicaceae

and Apiaceae) has also led to unsatisfactory generic limits Again the more general

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

19

problem is the use of one or a very few characters that have been weighted a priori to

structure classifications (see also Garciacutea et al 2009) although it is perhaps not

surprising that such an approach sometimes results in the delineation of taxa that have

indeed turned out to be quite distinct (cf eg Burtt Astragalus versus Oxytropis)

Taxa recognized here are monophyletic yet many are polythetic at the

morphological level that is they lack unique features characterizing (defining) all and

only members of that family they can be recognised phenetically only by the unique

combinations of characters that they posess This is the result of evolution any

synapomorphy characterizing a taxon may be lost or modified beyond easy recognition

in some of its members or the synapomorphy may appear to be identical to a feature

that has evolved in parallel in a quite unrelated plant That plant groups are polythetic is

almost as much a feature of monophyletic taxa as the paraphyletic taxa common in

evolutionary classifications Some families now include substantial variation as

phenetically distinct derived groups are placed in their proper phylogenetic position -

examples are Ericaceae which include the erstwhile Empetraceae a wind-pollinated

group that is florally very different and the overwhelmingly large-flowered and animal-

pollinated Plantaginaceae which now include several derived small-flowered aquatic

and wind-pollinated groups that were previously placed in separate families

Thus some families as delimited here may not be easy to recognize However

remember that detecting relationships - use whatever characters you can even if they

are not obvious - and naming a plant - focus on easy-to-see characters that may not

reflect relationships - are quite different problems Taxa although natural may not be

readily recognizable indeed it was in exactly this context that Lamarck worked out the

basic principles of writing dichotomous keys in 1778 (see Scharf 2007 for keys and the

like) Of course Lamarcks idea of nature was very different from ours - he thought that

there was some kind of continuum of form on which living organisms were to be

situated with no real gaps anywhere - but this meant that his genera (for example)

might well not be sharply distinct from each other even if each were part of the real

continuum that was life Lamarcks separation of the establishment of relationships and

groupings in a system and identification of the plants in that system have been widely

accepted (but cf eg Godfray amp Knapp 2004 many of the contributors to Heywood et

al 2007) Perhaps the best way of identifying plants at the family level is by well-made

multiple access keys as in Watson and Dallwitz (1992a onwards family limits there may

differ substantially from those adopted here) Multiple access keys free users from the

constraints of dichotomous keys in which particular characters are needed at each step

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

20

of the identification process before there can be further progress Instead those

characters that happen to be evident on a specimen can be used in whatever order is

convenient when linked to illustrations glossaries etc their power is enormous (see

Dallwitz et al 2000 [2006] for the principles underlying their construction and use)

Nevertheless dichotomous keys such as those of Hutchinson (1973) and Franz Thonner

(Geesink et al 1981) have their uses Of course taxa coming out adjacent in keys may

well not be at all related

(When identifying large numbers of plants even more efficient than either style of

identification and certainly lots more fun is sight identification Unless you have a

photographic memory you have to build up your knowledge of comparative plant

morphology - on which the ability to make accurate identifications depends - by

repeated observation When faced with an unknown plant I always look for leaf teeth

and stipules distinctive hair types especially stellate and T-shaped hairs glands and

punctations of any sort the presence of latex or other exudate and check leaf insertion

smelling crushed leaves can also be helpful In this context nodal anatomy can usually

be checked using a razor and a hand lens or even simply carefully examining leaf scars

The short paragraphs added after most families may help in confirming familial

identifications)

For the record and for the little that it is worth there are 4 orders and 13 families

of gymnosperms characterised on these pages and together they include some 82

genera and 947 species For angiosperms comparable figures are 56 orders 445

families 13208 genera and 261750 species (of which monocots include 11 orders 89

families 2759 genera and 52760 species) Note however that higher mathematics

was never my strong point and anyway these are pretty meaningless figures even for

species which many (but not all) might concede smacked slightly more of reality than

other taxa estimates range as high as 422000 (Govaerts 2001) Furthermore numbers

of genera and species change daily Nevertheless as emphasized here families are

useful in teaching we as a community can ensure that their limits remain largely stable

and by concentrating on relatively few of them one can gain some familiarity with much

of the worlds flora For summaries of the sizes of orders and families and of the general

arrangement followed in these pages see also the Statistics page itself

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

21

A PHYLOGENETIC TREE OF SEED PLANTS (THE MAIN TREE)

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

22

The Main Tree above is a conservative summary of well supported relationships

between the major clades of extant seed plants Reading - interpreting and

understanding - such a phylogenetic tree is not necessarily easy so reading a tree may

require a bit of practice OHara (eg 1988 1992) provides an accessible introduction to

thinking and talking about history in the context of looking at phylogenetic trees More

recently Krell and Cranston (2004) Crisp and Cook (2005) and others have emphasized

how careful one must be when interpreting and talking about evolution in ladderized

trees in particular and phylogenetic trees in general Gregory (2008) provides a

particularly detailed discussion of this problem In ladderized trees the smaller (in terms

of numbers of terminals) sister taxon is consistently shown on the same side at every

node thus the trees tend to be pectinate like the teeth of a comb however the

horizontal axis that is as a result so evident has no polarity or particular direction The

use of the adjective basal when discussing phylogenetic trees is especially dangerous

(see also Wojciechowski et al 2004 D Soltis et al 2005b) When I use the term and the

context is not otherwise clear I am referring to the pectinations at the base of a

ladderised tree Remember that when talking about sister taxa one can never be basal

to the other or older or younger although members of one can be more derived - in the

sense of having more apomorphies - than the other (but even then one has to be

careful) The words primitive and advanced and lower and higher should

practically never be used especially when talking about taxa Amborellaceae and

Pinaceae are sister to all other flowering plants and Pinales and are basal to the crown

groups of all other flowering plantsPinales respectively but that does not mean they

are thereby primitive Pinaceae in particular have numerous apomorphies The word

plesiomorphic is far less loaded than primitive and can be used to talk about

individual characters

Most trees here have been more or less ladderized as is the one above The

ladderization may be imperfect for example you can see that on the Main Tree

asterids with ten orders follow rosids and relatives which have seventeen although

this is in part because of the currently unresolved nature of relationships between the

main clades of core eudicots (this is likely to change soon) The Main Tree could have

been drawn with Amborella Acorus or a host of other taxa at the far right without

offending any relationships phylogenetic trees are like mobiles the only fixed points

being the nodes Despite the problems just mentioned pectination interpreted

carefully has its value and it is not an arbitrary process As one reads the terminals of a

pectinate tree from left to right adjacent terminals will be separated by apomorphies

that will cumulate in an appropriate fashion When thinking of a book or a herbarium

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

23

sequence (see Haston et al 2007) this is of some value Nymphaeales and

Austrobaileyales are here adjacent on the tree but they could be separated by

hundreds of families in the sequence merely by rotating only the node from which

Nymphaeales and their sister group (all other angiosperms minus about 75 species)

arise If adjacent in a book or herbarium or as in these summary pages then it is

relatively easy to relate their characters both apomorphic and plesiomorphic but if

separated by hundreds of pages or two floors in a large building then it is less easy to

get anything from the sequence (Since all orders in the pages of the main part of this

site are preceded by the apomorphies of all nodes immediately below them in the seed

plant phylogeny and because of the linkages that have been built in to the site this

problem is much less serious there there is no sequence) So the particular sequence of

all seed plants that is based on a phylogenetic classification but is to be used in herbaria

or any other place where taxa have to be arranged linearly as in this part of APweb

will be one that maximizes the number of taxa that are both successive branches of the

tree and placed successively in the sequence Since specimens are generally filed under

families the outline of a new family sequence for arranging herbaria and books can now

be suggested (Haston et al 2007) and such a standardized sequence will further help

teaching and learning about plants

ON THINKING ABOUT APOMORPHIES

Identifying apomorphies is important because understanding the evolution of

morphology in the broad sense - ie including anatomy chemistry etc - in the context

of diversification is one of our major goals For identifying apomorphies several

preconditions must be met One needs to have an accurate well supported phylogeny

one has to have examined the right taxa both from the point of view of morphology and

molecules one has to have coded the characters right (ie delimited states

appropriately) and one has to use an appropriate model of evolution when fitting the

variation to the tree (see eg Omland 1999 Stevens 2006b) If the distributions of

apomorphies below are compared with those in other studies (eg Turgeon et al 2001

Bremer et al 2001 Endress 2001 Albach et al 2001a Judd amp Olmstead 2004 D Soltis

et al 2005b Zhang et al 2006 Judd et al 2007) the reader may find a number of

differences Although I have integrated such studies as far as is possible there are five

reasons that link to the preconditions just mentioned why there may be differences

where features are placed on trees

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

24

1 I may not have found all the information about a particular character there may be

disagreement over its interpretation or I have added information

2 The sampling of nearly all molecular studies is very incomplete (see Salisbury amp Kim

2001 for problems caused by sampling) indeed it is commonplace to decry the

incompleteness of molecular sampling But not only is the sampling in molecular studies

often less than we might wish that of the morphological and chemical characters whose

evolution we are interested in understanding is also often very poor So for many

anatomical chemical and embryological characters that are confidently said to

characterise families and other groups we all too often have no idea if those characters

are applicable to the whole clade or just to a subgroup within that clade However as

our sampling improves we can locate changes on the tree more precisely if in

unexpected positions Thus is is sometimes suggested that Ericaceae have ellagic acid

(Soltis et al 2005b) in fact the little evidence we have indicates that only a small clade

of some 80 species has ellagic acid and the rest of the family - itself alone about one

third of all Ericales - does not Similarly Ericaceae are noted for their

ectendomycorrhizal associations but Enkianthus sister to all other Ericaceae

apparently lacks these (Abe 2005) Thus these mycorrhizae and their associated

distinctive hair roots are unlikely to be apomorphies of Ericaceae - however it appears

that other families near Ericaceae may also have ectendomycorrhizae (see Asai 1934)

3 I am fitting characters to a very conservative tree with many polytomies although the

nodes that are utilised are for the most part strongly supported polytomies make the

optimisation of characters that is the assigment of character state change to a

particular node on the tree notably difficult (eg Madison amp Madison 2002) In nearly

all studies of the evolution of characters distributions of characters are optimised on a

more or less fully resolved tree and the construction of supertrees may yield yet more

detailed hypotheses of relationships (for literature on supertrees see Cotton amp

Wilkinson 2007 2008) Of course some nodes on such fully resolved trees andor

supertrees may have little support and optimisations of characters on such trees may

carry correspondingly little conviction Even parts of some of the trees used here have

poor support eg relationships within aquatic Alismatales etc although I indicate

when this is the case

4 Exactly how one goes about optimising a character on a tree is critically important

Even using simple parsimony optimisations (ACCTRAN or DELTRAN ACCelerated

TRANsitions or DELayed TRANsitions) the position of synapomorphies on trees - and

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

25

hence our ideas of evolution - may differ and this is still more true if one uses maximum

likelihood or Bayesian methods and makes apparently reasonable suggestions about

weighting gains over losses (eg Donoghue amp Ackerley 1996 Cunningham et al 1998

Omland 1997 1999 Ree amp Donoghue 1999 Polly 2001 Webster amp Purvis 2001

Ronquist 2004 Crisp amp Cook 2005) Sannier et al (2007) show how in in Arecaceae that

where on a phylogenetic tree one might peg changes in microsporogenesis will depend

on the methods one uses to do this and Pedersen et al (2007) discuss the sometimes

very substantial effect of node support on the posterior probabilities of ancestral

character states Here I use parsimony optimization not always as explicit as it might be

but I have often indicated where there are particularly important uncertainties as to the

positions of particular character changes on the tree

5 Finally although I have paid quite a lot of attention to the delimitation of the

character states that make up all the characterizations I have not spent enough time on

this critical operation If we are interested in understanding evolution then fitting the

basic variation - not character states - to a tree in principle allows greater flexibility in

understanding morphology in the context of local phylogenies (see also Stevens 2000

Endress 2005c) However many character states used here are delimited globally that

is they are circumscribed in the context of the variation shown by individual characters

across all angiosperms andor in the context of classic ideas of character evolution

Character states often have arbitrary limits and serve best to communicate

information whether they are in fact suitable for either phylogenetic analysis or

understanding evolution are separate issues Studies have rather unsurprisingly

perhaps but importantly shown that dividing the one character into different sets of

states may yield differing ideas of evolution of that character (eg Lamb Frye amp Kron

2003 Hibbett 2004) When looking at trees on which character states are optimised

one should bear in mind the problems surrounding the delimitation of states (eg

Stevens 2000 2006b) and the danger of using pollen or other types - constructs

based on many characters that vary independently but which effectively get lost in

these types thus Blackmore et al (2009) decomposed the pollen types in Asteraceae

into 52 characters

I use Remanes three main criteria of homology or better similarity when

determining the basic similarity of structures on different organisms (see Remane 1952)

These criteria are special properties position and intermediates Special properties

include anatomical or chemical characters gene expression data etc Position refers

to the position of an organ with respect to landmarks on the plant Although plants are

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

26

plants and landmarks are not as fixed as one might like trying to work out the relative

positions of parts is a good way of understanding morphology so long as one does not

try to out-do Procrustes Intermediates include intermediates found on the same

plant or on different plants Intermediates may observed during development or be

adult structures Thus one may find changes during development which suggest what

the nature of a particular structure is and very different stuctures on different plants

may turn out to be similar early in development Or one can simply compare different

taxa and see that structures that appear to be very different when comparing two

different taxa can be linked morphologically by looking at other taxa

Nevertheless the use of these criteria may not yield an unambiguous answer as to

what a structure is even given a solid phylogeny and an improved understanding of

development (see Jaramillo amp Kramer 2007 for a useful discussion) As Endress (2005c)

observed a number of features - position function development shape anatomy

histology gene activity and relationships to other taxa that clearly have petals - can be

used to distinguish a petal (for example) from other floral structures if a petal does not

have one of these features is it thereby not a petal Thus Maturen et al (2005) recently

found that floral organ diversity genes (B and C) were expressed in the large white

inflorescence bracts of Cornus (see also Costa et al 2005) Peney et al (2005) noted that

not all monosulcate pollen grains in monocots have the same developmental pathway

and that as a result such pollen might not have the same ancestral state Reeves and

Olmstead (2003) suggested that the genetic mechanisms causing monosymmetry in

Lamiales and Solanales were different and Serna and Martin (2006) described similar

problems with the development of hairs in Arabidopsis when compared with that of

hairs in Antirrhinum and Solanaceae Indeed as one perhaps might expect delimitation

of states and characters does not necessarily become easier with increasing knowledge

of development etc Thus Buzgo et al (2004) Matthews and Endress (2005) and others

have shown how hard it can be to distinguish between eg prophylls and other floral

structures as their behaviour is studied during the course of the development of the

flower Similarly at what concentration is a particular secondary metabolite deemed to

be present (Waterman (2007)

The validity of the approach used here that of fitting morphological variation to a

largely molecular-based tree may be questioned However I think it rather unlikely that

well-supported molecular branches will be overturned by morphological data Indeed

analyses of morphological data alone do provide support for many of the clades evident

in molecule-only analyses and analyses with morphological and molecular data

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

27

together may lead to increased support for clades (eg Hufford 1992 Nandi et al 1998

[but adding morphological data reduces support for a number of critical clades too]

Doyle amp Endress 2000 however in none of these papers is the use of morphology

without ambiguity) It is unfortunately clear that the use of morphology alone may not

suggest problems in the phylogenetic placement of taxa that later turn out to have been

wrongly included (eg Zhang et al 1992) Thus I have been wary of putting much weight

on clades that have only morphological support but note that I have treated molecule-

based clades with low bootstrap or jacknife support values (esp below 70) or low

posterior probabilies (below 095) likewise Although I may have been mistaken in

placing so much emphasis on molecular data in terms of providing the basic

phylogenetic framework for angiosperms morphological and molecular data are only

very rarely in irreconcilably strong conflict There are indeed a few places where the

conflict seems extreme These include the relative positions of the Monimiaceae and

Hernandiaceae (Laurales) the position of Hanguanaceae (Commelinales [as here] or

Zingiberales) and of Triplostegia (is it in Dipsacaceae or Valerianaceae - see

Dipsacales) Fossils are unlikely to affect the topologies of the trees presented here but

see below for their importance in understanding morphological evolution in general and

the evolution of angiosperms in particular Nevertheless some largely reject the idea

that trees based on molecular data alone can recover phylogenetic relationships

especially when branching points are old and prefer to used trees based on analysis of

morphological data including those taken from fossils (Hilton amp Bateman 2006 Farjon

2007)

All in all however the extent of the congruence between morphological and

molecular data is impressive and heartening and many clades can be characterised

morphologically It seemed in 1998 that there were no unambiguous morphological

synapomorphies for angiosperm orders (K Bremer 2000) and this is still true if by

unambiguous is meant non-homoplasious However many orders and other clades

have synapomorphies even if these may be indistinguishable at least at the current

level of morphological and developmental knowledge from parallel occurences

elsewhere As our morphological knowledge increases so too does the number of

apomorphies

Endress and Matthews (2006a) emphasize the importance of tendencies and

developmental constraints when thinking about characters of clades within the rosids -

many characters or character combinations occur in a rather sporadic fashion within

the clade and apparently notably less frequently outside it There are a number of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

28

examples of tendencies here and in the individual order pages such as the features

enclosed by parentheses in groups above the level of family eg see (cuticular waxes

as aggregated rodlets) for the commelinids and the discussion of the distribution of

polyandry that is flowers with numerous stamens in the asterid I + II groups etc

Tendencies may involve more than single characters When thinking of the

associations of features like integument thickness and vascularization of the integument

(Endress amp Matthews 2006a) one can imagine fairly simple developmental

preconditions being operative As Endress and Matthews (2006a) note it is difficult to

imagine vascular tissue developing in an integument only two cells thick Changes here

would be loosely correlated if morphologically linked However the strongly correlated

changes noted by Givnish et al (2005) are ecologically linked but are presumably

morphologicallydevelopmentally independent When there seem to be characters

evolving more or less together tests can be carried out to see if the changes are

concentrated on certain branches of the tree (eg Maddison 1990 Sanderson 1991

Maddison amp Maddison 2000) Such tests have rarely been carried out at higher levels in

angiosperms

One of the most striking examples of a tendency is the distribution of N-fixation

restricted as it is to a monophyletic group of four clades although it has arisen seven (or

perhaps several more) times independently within the N-fixing clade and several

members of two quite different kinds of bacteria are involved (eg Clawson et al 2004

Elliott et al 2007 Sprent amp James 2007) Other examples of work that bears on the issue

of tendencies include the findings that flowers of polysymmetrical Arabidopsis have

genes like TCP1 that are expressed asymmetrically during early development and TCP1

is a probable orthologue of the well-known CYC gene of Antirrhinum that is involved in

the development of monosymmetric slowers there (Cubas et al 2001 Costa et al 2005

etc) Parallelism might build on this underlying morphologically cryptic monosymmetry

even if details of the genetic mechanisms causing the monosymmetry evident in

particular groups may be different (Reeves amp Olmstead 2003 Cubas 2004 see above)

Similarly there have been several recent suggestions that the capability to synthesise a

particular metabolite may be switched off but not lost and so can sometimes be

reacquired (eg Wink amp Witte 1983 Wink 2003 Liscombe et al 2005 Larsson 2007

Waterman 2007) Hence perhaps the rather spotty distribution of many secondary

metabolites like ellagic acid the indole alkaloid camptothecin iridoids etc when

considered in the context of phylogenies As a non-botanical example - but a rather nice

one - Salwini-Plawen and Mayr (1961) suggested some time ago that there has been

considerable parallelism (40-65 or more independent origins) in the evolution of eyes in

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

29

metazoans However Pax 6 seems to be a master control gene that is involved in eye

formation perhaps throughout all bilateralians (eg Gehring amp Ikeo 1999 Erwin amp

Davidson 2002) which changes the nature of the problem Note however that in

general our invocation of evolutionary tendencies to explain the patterns of

distributions of characters is really an admission that we do not understand those

patterns

As we find out more about variation we will probably find fewer and fewer features

constant throughout a group Most unqualified statements of presence and absence

should properly be qualified as usually present or usually absent if one is thinking of

the characterisations on the order pages as encompassing the total variation within a

clade Thus Pistia alone among monocots as so far known has sieve tube plastids with

starch grains not protein crystals However this does not affect the fact that sieve tube

plastids with cuneate protein crystals and lacking starch grains are an apomorphy for

monocots Furthermore fossils in a number of cases suggest character combinations

unknown in extant taxa as may be seen in the discussions of Fagaceae Platanaceae

Iteaceae Calycanthaceae etc Confusing the issue there may be questions as to where

exactly on the tree a particular fossil is to be placed (eg see Nymphaeaceae

Calycanthaceae Archaefructus etc)

To summarize given our current understandings of both phylogenies and

characters evolution of some characters in which we are interested seems very labile

(see eg D Soltis et al 2005b Endress and Matthews 2006a Stevens 2006b Ekman et

al 2008 for a good recent study) and I have been cautious when talking about character

evolution Much effort must continue to be spent in summarizing characters of well-

established clades at all levels providing features by which they may be recognized and

signaling synapomorphies Remember that (1) the basic morphological anatomical and

chemical knowledge of many critical taxa is woefully incomplete (2) different

assumptions about character evolution may greatly affect the position of

synapomorphies on trees (3) in many cases relationships within and between many

groups are too uncertain at present to worry very much about synapomorphies and

(4) we must be clear about what we do and do not not know As mentioned above I

have indicated in a number of places where there are particularly important

uncertainties as to where characters should be placed on the tree Nevertheless it is a

relatively easy matter to update notes such as these and it can be a simple matter to

incorporate new data on characters that have never before been considered in the

context of a tree Much basic - and unfortunately perhaps unfashionable - work must be

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

30

carried to clarify the distribution of morphological anatomical and chemical characters

for examples of what can to be done see the work of P K Endress and collaborators

(floral morphology and development) and S R Jensen and collaborators (iridoids) But

acquiring information about nodal anatomy cell and tissue distribution nodal anatomy

and the like is not difficult at all and should be generally encouraged

But all this is merely a necesary prelude to the understanding of evolution For this

one needs to know a lot more including dating the phylogeny understanding the

function(s) of characters factoring in relevant aspects of the palaeoenvironment in

which an apomorphy first appeared etc

SUMMARY OF THE SYSTEM

Below is a formalised summary of the relationships within orders of the families of seed

plants There are a few families that are not recognised even as options in APG II and

vice versa as well as a few extra orders The families to be recognised in the most

recent edition of Mabberleys The Plant Book (Mabberley 2008) are also largely

consistent with those below But all differences are trivial and will - I hope - eventually

disappear

Square brackets - [] - enclose clades the plus sign - + - designates sister taxa a

comma - - denotes part of a polytomy and quotation marks - - denotes a

paraphyletic group

SEED PLANTS

GYMNOSPERMS

Cycadales

Cycadaceae + Zamiaceae

Ginkgoales

Ginkgoaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

31

Pinales

Pinaceae [[Araucariaceae [Podocarpaceae + Phyllocladaceae]] [Sciadopityaceae

[Taxaceae + Cupressaceae]]]

Gnetales

Ephedraceae [Gnetaceae + Welwitschiaceae]

ANGIOSPERMSFLOWERING PLANTS

Amborellales

Amborellaceae

Nymphaeales

Hydatellaceae [Cabombaceae + Nymphaeaceae]

Austrobaileyales

Austrobaileyaceae [Schisandraceae + Trimeniaceae]

Chloranthales

Chloranthaceae

MAGNOLIIDS

Magnoliales

Myristicaceae [Magnoliaceae [[Himantandraceae + Degeneriaceae] [Eupomatiaceae +

Annonaceae]]]

Laurales

Calycanthaceae [[Siparunaceae [Gomortegaceae + Atherospermataceae]] [Monimiaceae

[Hernandiaceae + Lauraceae]]]

Canellales

Canellaceae + Winteraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

32

Piperales

[Hydnoraceae + Aristolochiaceae] [Piperaceae + Saururaceae]

MONOCOTS

Acorales

Acoraceae

Alismatales

Araceae [Tofieldiaceae [[Alismataceae [Hydrocharitaceae + Butomaceae]]

[Scheuchzeriaceae [Aponogetonaceae [Juncaginaceae [Maundiaceae [[Posidoniaceae

[Ruppiaceae + Cymodoceaceae]] [Zosteraceae + Potamogetonaceae]]]]]]]

Petrosaviales

Petrosaviaceae

Dioscoreales

Nartheciaceae [[Taccaceae + Thismiaceae] [Burmanniaceae + Dioscoreaceae]]

Pandanales

Velloziaceae Triuridaceae Stemonaceae [Pandanaceae + Cyclanthaceae]

Liliales

Corsiaceae [Campynemataceae [Petermanniaceae [Colchicaceae + Alstroemeriaceae]]

Melanthiaceae [[Philesiaceae + Rhipogonaceae] [Smilacaceae + Liliaceae]]]

Asparagales

Orchidaceae [[Boryaceae [Blandfordiaceae [Lanariaceae [Asteliaceae + Hypoxidaceae]]]]

[[Ixioliriaceae + Tecophilaeaceae] [Doryanthaceae [Iridaceae [Xeronemataceae

[Xanthorrhoeaceae [Alliaceae + Asparagaceae]]]]]]]

COMMELINIDS

Unplaced

Dasypogonaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

33

Arecales

Arecaceae

Poales

[Typhaceae + Bromeliaceae] [Rapateaceae [[Thurniaceae [Juncaceae + Cyperaceae]]

[[Mayacaceae [Eriocaulaceae + Xyridaceae]] [Flagellariaceae [Anarthriaceae

[Centrolepidaceae + Restionaceae]]] [Joinvilleaceae [Ecdeiocoleaceae + Poaceae]]]]]]]

Commelinales

[Commelinaceae + Hanguanaceae] [Philydraceae [Haemodoraceae + Pontederiaceae]]

Zingiberales

Musaceae [Strelitziaceae + Lowiaceae] Heliconiaceae [[Cannaceae + Marantaceae]

[Costaceae + Zingiberaceae]]

Ceratophyllales

Ceratophyllaceae

EUDICOTS

Ranunculales

Eupteleaceae [Papaveraceae [[[Lardizabalaceae + Circaeasteraceae] [Menispermaceae

[Berberidaceae + Ranunculaceae]]]]

Sabiales

Sabiaceae

Proteales

Nelumbonaceae [Platanaceae + Proteaceae]

Trochodendrales

Trochodendraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

34

Buxales

Haptanthaceae [Buxaceae + Didymelaceae]

CORE EUDICOTS

Gunnerales

Gunneraceae + Myrothamnaceae

Dilleniales

Dilleniaceae

Saxifragales

Peridiscaceae [[Paeoniaceae [Altingiaceae [Hamamelidaceae [Cercidiphyllaceae +

Daphniphyllaceae]]]] [[Crassulaceae [Aphanopetalaceae [Tetracarpaeaceae

[Penthoraceae + Haloragaceae]]]] [Iteaceae [Grossulariaceae + Saxifragaceae]]]]

Cynomoriaceae unplaced

Vitales

Vitaceae

ROSIDS

FABIDROSID I

Zygophyllales

Krameriaceae + Zygophyllaceae

Celastrales

Lepidobotryaceae + Celastraceae

Oxalidales

Huaceae [[Connaraceae + Oxalidaceae] [Cunoniaceae [Elaeocarpaceae [Brunelliaceae +

Cephalotaceae]]]]

Malpighiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

35

[Achariaceae [Goupiaceae [Violaceae + Passifloraceae] [Lacistemataceae Salicaceae]]

[Lophopyxidaceae Putranjivaceae] [Ctenolophonaceae [Erythroxylaceae +

Rhizophoraceae]] Linaceae Ixonanthaceae Humiriaceae Irvingiaceae

Centroplacaceae Pandaceae [[Bonnetiaceae + Clusiaceae] [Calophyllaceae

[Hypericaceae + Podostemaceae]]] [Malpighiaceae + Elatinaceae] Ochnaceae

[Peraceae [Rafflesiaceae + Euphorbiaceae]] [Phyllanthaceae + Picrodendraceae]

[Balanopaceae [[Trigoniaceae + Dichapetalaceae] [Chrysobalanaceae +

Euphroniaceae]]] Caryocaraceae

N-FIXING CLADE

Fabales

Quillajaceae [Fabaceae [Polygalaceae + Surianaceae]]

Rosales

Rosaceae [[Barbeyaceae [Elaeagnaceae Dirachmaceae Rhamnaceae]] [Ulmaceae

[Cannabaceae [Moraceae + Urticaceae]]]]

Cucurbitales

Anisophylleaceae [[Corynocarpaceae + Coriariaceae] [Cucurbitaceae [Tetramelaceae

[Datiscaceae + Begoniaceae]]]] Apodanthaceae

Fagales

Nothofagaceae [Fagaceae [[Myricaceae + Juglandaceae] [Casuarinaceae

[Ticodendraceae + Betulaceae]]]]

MALVIDROSID II

pgtGeraniales

Geraniaceae [[Melianthaceae + Francoaceae] [Vivianaceae + Ledocarpaceae]]

Myrtales

Combretaceae [[Onagraceae + Lythraceae] [[Vochysiaceae + Myrtaceae]

[Melastomataceae [Crypteroniaceae [Alzateaceae + Penaeaceae]]]]]

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

36

Crossosomatales

[Staphyleaceae [Guamatelaceae [Crossosomataceae + Stachyuraceae]]] [Aphloiaceae

[Geissolomataceae + Strasburgeriaceae]]

Picramniales

Picramniaceae

Sapindales

Biebersteiniaceae Nitrariaceae [Kirkiaceae [Anacardiaceae + Burseraceae]]

[Sapindaceae [Simaroubaceae Rutaceae Meliaceae]]

Huerteales

Gerradinaceae [Dipentodontaceae + Tapisciaceae]

Malvales

Neuradaceae [Thymelaeaceae [Sphaerosepalaceae Bixaceae [Cistaceae

[Sarcolaenaceae + Dipterocarpaceae]] [Cytinaceae + Muntingiaceae] Malvaceae]]

Brassicales

[Akaniaceae + Tropaeolaceae] [[Moringaceae + Caricaceae] [Setchellanthaceae

[Limnanthaceae [[Koeberliniaceae [Bataceae + Salvadoraceae]] [Emblingiaceae

[Pentadiplandraceae [Gyrostemonaceae Resedaceae Stixis and relatives] Tovariaceae

[Capparaceae [Cleomaceae + Brassicaceae]]]]]]]]

Berberidopsidales

Aextoxicaceae + Berberidopsidaceae

Santalales

Erythropalaceae Olacaceae [[Loranthaceae [Misodendraceae + Schoepfiaceae]]

[Opiliaceae + Santalaceae]] Balanophoraceae unplaced

Caryophyllales

[[Droseraceae [Nepenthaceae [Drosophyllaceae [Ancistrocladaceae +

Dioncophylleaceae]]]]] [[Frankeniaceae + Tamaricaceae] [Polygonaceae +

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

37

Plumbaginaceae]]] [Rhabdodendraceae [Simmondsiaceae [[Asteropeiaceae +

Physenaceae] [[Caryophyllaceae [Achatocarpaceae + Amaranthaceae]]

[Stegnospermataceae [Limeaceae [[Lophocarpaceae [Barbeuiaceae [Aizoaceae

[Sarcobataceae Phytolaccaceae Nyctaginaceae]]]] [Molluginaceae [Halophytaceae

Montiaceae Basellaceae Didiereaceae [Talinaceae [Portulacaceae [Anacampseros etc +

Cactaceae]]]]]]]]]]]]]

ASTERIDS

Cornales

[Cornaceae + Nyssaceae] [Hydrangeaceae + Loasaceae] Hydrostachyaceae [Curtisiaceae

+ Grubbiaceae]

Ericales

[Balsaminaceae [Marcgraviaceae + Tetrameristaceae]] [[Polemoniaceae +

Fouquieraceae] Lecythidaceae [[Sladeniaceae + Pentaphylacaceae] [Sapotaceae

[Ebenaceae [Maesaceae [Theophrastaceae [Myrsinaceae + Primulaceae]]]]]

[Mitrastemonaceae Theaceae [Symplocaceae [Styracaceae + Diapensiaceae]]

[[Sarraceniaceae [Roridulaceae + Actinidiaceae] [Clethraceae [Cyrillaceae +

Ericaceae]]]]]]

LAMIIDASTERID I

Unplaced

Metteniusaceae Oncothecaceae Icacinaceae and some unplaced ex-Icacinaceae

Garryales

Garryaceae + Eucommiaceae

Unplaced

Boraginaceae Vahliaceae

Gentianales

Rubiaceae [Gentianaceae [Loganiaceae [Gelsemiaceae + Apocynaceae]]]

Lamiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

38

Plocospermataceae [Oleaceae [Tetrachondraceae [Calceolariaceae

[Gesneriaceae Peltanthera [Pedaliaceae Carlemanniaceae Martyniaceae Stilbaceae

Plantaginaceae Linderniaceae Bignoniaceae Verbenaceae Lamiaceae Paulowniaceae

Schlegeliaceae Thomandersiaceae Phrymaceae Rehmannia Acanthaceae

Scrophulariaceae Orobanchaceae Byblidaceae Lentibulariaceae]]]]]

Solanales

[Montiniaceae [Sphenocleaceae + Hydroleaceae]] [Convolvulaceae + Solanaceae]

CAMPANULIDASTERID II

Aquifoliales

[[Cardiopteridaceae + Stemonuraceae] [Aquifoliaceae [Helwingiaceae +

Phyllonomaceae]]

Asterales

[Rousseaceae + Campanulaceae] Pentaphragmataceae [[Alseuosmiaceae [Phellinaceae

+ Argophyllaceae]] Stylidiaceae [Menyanthaceae [Goodeniaceae [Calyceraceae +

Asteraceae]]]]

Escalloniales

Escalloniaceae

Bruniales

[Bruniaceae + Columelliaceae]

Apiales

Pennantiaceae [Torricelliaceae [Griseliniaceae [Pittosporaceae [Araliaceae

[Myodocarpaceae + Apiaceae]]]]]

Paracryphiales

Paracryphiaceae

Dipsacales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

39

Adoxaceae [Diervillaceae [Caprifoliaceae [Linnaeaceae [Morinaceae [Dipsacaceae +

Valerianaceae]]]]]

LINKS TO ORDERS AND FAMILIES

[Back to Top]

MAIN GROUPINGS

Asterids commelinids core eudicots asterid 1 asterid

2 eudicots gymnosperms Magnoliophyta monocots N-fixing claderosids seed plants

ALPHABETICAL LISTING OF ALL ORDINAL NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | L | M | N | O | P | Q | R | S | T | U | V | W | X | Z

Abietales Acanthales Acerales Acorales Actinidiales Actinostrobales Adoxales Aescul

ales Agavales Aizoales AkanialesAlismatales Alliales Alseuosmiales Alstroemeriales

Altingiales Amaranthales Amaryllidales Amborellales AmbrosialesAmmiales Amomal

es Ancistrocladales Anisophylleales Annonales Anthobolales Apiales Apocynales Apo

nogetonalesAquifoliales Arales Araliales Aralidiales Araucariales Arecales Aristolochi

ales Athrotaxidales Asarales AsclepiadalesAsparagales Asphodelales Asparagales Ast

eliales Atriplicales Aucubales Austrobaileyales Avenales

Balanitales Balanopales Balanophorales Balsaminales Barbeyales Barclayales Batales

Begoniales BerberidalesBerberidopsidales Betulales Biebersteiniales Bignoniales Bix

ales Boraginales Brassicales Brexiales Bromeliales BrunialesBrunoniales Burmannial

es Burserales Butomales Buxales Byblidales

Cactales Callitrichales Calycanthales Calycerales Campanulales Campynematales Can

ellales Cannales CapparalesCaprifoliales Cardiopteridales Carduales Caricales Carle

manniales Caryophyllales Cassiales Casuarinales CelastralesCentrolepidales Cephalot

ales Cephalotaxales Ceratophyllales Cercidiphyllales Chenopodiales Chironiales Chlor

anthalesChrysobalanales Cinchonales Circaeasterales Cistales Citrales Cocosales Col

chicales Columelliales CombretalesCommelinales Connarales Convolvulales Coriarial

es Cornales Corylales Corynocarpales Crassulales CrossosomatalesCucurbitales Cunn

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

40

inghamiales Cunoniales Cupressales Cycadales Cyclanthales Cymodoceales Cynarales

CynomorialesCyperales Cyrillales Cytinales

Daphnales Daphniphyllales Dasypogonales Datiscales Degeneriales Desfontainiales D

iapensiales Didymelales DillenialesDioales Dioncophyllales Dioscoreales Diospyrales

Dipentodontales Dipsacales Droserales

Ebenales Echiales Elaeagnales Elaeocarpales Elatinales Elodeales Emmotales Empetr

ales Ephedrales Ericales EriocaulalesErythropalaless Escalloniales Eucommiales Euph

orbiales Eupomatiales Eupteleales Euryalaless

Fabales Fagales Falcatifoliales Ficales Flacourtiales Flagellariales Fouquieriales Franc

oales Frangulales

Galiales Garryales Geissolomatales Gentianales Geraniales Gesneriales Ginkgoales G

laucidiales Globulariales GnetalesGoodeniales Greyiales Griseliniales Grossulariales

Grubbiales Gunnerales Gyrocarpales Gyrostemonales

Haemodorales Haloragales Hamamelidales Hanguanales Heisteriales Helleborales He

lwingiales Himantandrales HippuridalesHomaliales Hortensiales Huales Huerteales

Hydatellales Hydnorales Hydrangeales Hydrastidales HydrocharitalesHydropeltidales

Hydrostachyales Hypericales Hypoxidales

Icacinales Illiciales Iridales Irvingales Iteales Ixiales

Jasminales Juglandales Julianiales Juncaginales Juncales

Lacistematales Lactoridales Lamiales Lardizabalales Laurales Lecythidales Ledocarpal

es Leitneriales LentibularialesLigustrales Liliales Limnanthales Linales Liriales Loasal

es Lobeliales Loganiales Lonicerales Loranthales LowialesLythrales

Magnoliales Malpighiales Malvales Marathrales Marcgraviales Mayacales Medusagy

nales Medusandrales MelanthialesMelastomatales Meliales Melianthales Meliosmal

es Menispermales Menyanthales Metteniusales MitrastemonalesMiyoshiales Moni

miales Moringales Musales Myricales Myristicales Myrothamnales Myrsinales Myrta

les

Najadales Nandinales Narcissales Nartheciales Nelumbonales Nepenthales Neuradal

es Nitrariales Nolanales NothofagalesNyctaginales Nymphaeales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

41

Ochnales Oenotherales Olacales Oleales Onagrales Oncothecales Opuntiales Orchid

ales Oxalidales

Saxifragales Pandales Pandanales Papaverales Paracryphiales Parasitaxales Paridales

Parnassiales PassifloralesPenaeales Pennantiales Pentaphragmatales Peridiscales P

etiveriales Petrosaviales Phellinales Philydrales PhyllanthalesPhysenales Phytolaccal

es Picramniales Pinales Pinguiculales Piperales Pittosporales Plantaginales Platanale

sPlumbaginales Poales Podocarpales Podophyllales Podostemales Polemoniales Pol

ygalales Polygonales PontederialesPortulacales Posidoniales Potamogetonales Primu

lales Proteales

Quercales Quillajales Quintiniales

Rafflesiales Ranunculales Rapateales Resedales Restionales Rhabdodendrales Rhamn

ales Rhinanthales RhizophoralesRhodorales Rhoipteleales Roridulales Rosales Rouss

eales Rubiales Ruppiales Rutales

Sabiales Salicales Salvadorales Samolales Samydales Sanguisorbales Santalales Sapin

dales Sapotales SarracenialesSaxegotheales Saxifragales Scheuchzeriales Sciadopitya

les Scleranthales Scrophulariales Scyphostegiales SedalesSilenales Simmondsiales S

milacales Solanales Sphenocleales Sphenostemonales Stangeriales Stellariales Stemo

nalesStilbales Stylidiales Styracales Surianales

Taccales Tamales Tamaricales Taxales Taxodiales Tecophilaeales Terebinthales Tern

stroemiales Theales TheligonalesThymelaeales Tiliales Tofieldiales Torricelliales Tov

ariales Tribelales Trilliales Trimeniales Triuridales TrochodendralesTropaeolales Turn

erales Typhales

Ulmales Urticales

Vacciniales Vahliales Vallisneriales Velloziales Veratrales Verbenales Viburnales Vinc

ales Violales Viscales VitalesVochysiales

Welwitschiales Winterales

Xanthorrhoeales Ximeniales Xyridales

Zamiales Zingiberales Zosterales Zygophyllales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

42

ALPHABETICAL LISTING OF ALL FAMILY NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | K | L | M | N | O | P | Q | R | S | T | U | V | W | X | Y

|Z

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raceae Aragoaceae Araliaceae Aralidiaceaee Araucariaceae Arbutaceae Arceuthidace

aeArceuthobiaceae Arctostaphylaceae Arctotidaceae Ardisiaceae Arecaceae Argoph

yllaceae Arisaraceae AristolochiaceaeAristoteliaceae Arjonaceae Armeriaceae Artem

isiaceae Arthrotaxidaceae Artocarpaceae Arundinaceae ArundinellaceaeAsaraceae A

sclepiadaceae Ascyraceaee Asiraceae Asparagaceae Asperulaceae Asphodelaceae As

pidistraceaeeAsteliaceae Asteraceae Asteranthaceae Asteropeiaceae Astragalaceae

Astrocarpaceae AthanasiaceaeAtherospermataceae Atriplicaceae Atropaceae Aucuba

ceae Aurantiaceae Austrobaileyaceae Austrotaxaceae AvenaceaeAverrhoaceae Avet

raceae Avicenniaceae Azaleaceae Azimaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Balanitaceae Balanopaceae Balanophoraceae Balsameaceae Balsaminaceae Bambusa

ceae Barbaceniaceae BarbeuiaceaeBarbeyaceae Barclayaceae Barringtoniaceae Base

llaceae Bataceae Baueraceae Bauhiniaceae Baxteriaceae BegoniaceaeBehniaceae B

elangeraceae Belloniaceae Bembiciaceae Berberidaceae Berberidopsidaceae Berryac

eaee BersamaceaeBertyaceae Berzeliaceae Besleriaceae Betaceae Betulaceae Biebe

rsteiniaceae Bifariaceae Bignoniaceae BischofiaceaeBixaceae Blakeaceae Blandfordia

ceae Blattiaceae Blepharocaryaceae Blitaceae Blyxaceae Boerlagellaceae Bolivaracea

eBombacaceae Bonnetiaceae Bontiaceae Boopidaceae Boraginaceae Borassaceae B

oroniaceae BoryaceaeBotryodendraceae Bougainvilleaceae Boweniaceae Brassicacea

e Brachycaulaceae Bretschneideraceae BrexiaceaeBromeliaceae Brownlowiaceae Br

unelliaceae Bruniaceae Brunoniaceae Brunsvigiaceae Bryoniaceae BuchneraceaeBuci

daceae Buddlejaceae Buglossaceae Bulbocodiaceae Bumeliaceae Bupleuraceae Burc

hardiaceae BurmanniaceaeBurseraceae Butneriaceae Butomaceae Buxaceae Byblida

ceae Byttneriaceae

Cabombaceae Cacaoaceae Cactaceae Caesalpiniaceae Caladiaceae Calamaceae Calce

olariaceae CalectasiaceaeCalendulaceae Callaceae Callicomaceae Calligonaceae Calli

traceae Callitrichaceae Calochortaceae CalophyllaceaeCalthaceae Calycanthaceae Ca

lyceraceae Cambogiaceae Camelliaceae Campanulaceae CampynemataceaeCanacom

yricaceae Canellaceae Cannabaceae Cannaceae Canopodaceae Canotiaceae Cansjera

ceae CapparaceaeCaprariaceae Caprifoliaceae Cardiopteridaceae Carduaceae Carica

ceae Carissaceae Carlemanniaceae CarpinaceaeCarpodetaceae Cartonemataceae Ca

ryocaraceae Caryophyllaceae Casearieaceae Cassiaceae CassipoureaceaeCassythacea

e Castaneaceae Castelaceae Casuarinaceae Catesbaeaceae Cathedraceae Cecropiace

ae CedraceaeCedrelaceae Celastraceae Celosiaceae Celtidaceae Centaureaceae Cen

trolepidaceae Centroplacaceae CepaceaeCephalanthaceae Cephalotaceae Cephalota

xaceae Cerastiaceae Ceratoniaceae Ceratophyllaceae CerberaceaeCercidiphyllaceae

Cercocarpaceae Cercodiaceae Cereaceae Cerinthaceae Ceroxylaceae Cestraceae Cev

alliaceaeChailletiaceae Chamaedoraceae Chamaemoraceae Chamelauciaceae Chaun

ochitonaceae Chelidoniaceae ChelonaceaeChenopodiaceae Chimonanthaceae Chingi

thamnaceae Chionographidaceae Chiranthodendraceae ChironiaceaeChlaenaceae Ch

loanthaceae Chloranthaceae Chloridaceae Chlorogalaceae Chrysobalanaceae Cicerac

eae CichoriaceaeCimicifugaceae Cinchoniaceae Circaeaceae Circaeasteraceae Cissac

eae Cistaceae Citraceae Clematidaceae CleomaceaeClethraceae Clusiaceae Cneorac

eae Cnestidaceae Cnicaceae Cobaeaceae Cochlospermaceae Cocosaceae Coffeaceae

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Colchicaceae Coleogynaceae Columelliaceae Combretaceae Commelinaceae Comocl

adiaceae Compositae CompsoaceaeConantheraceae Connaraceae Conostylidaceae C

onvallariaceae Convolvulaceae Cordiaceae CoreopsidaceaeCoriandraceae Coriariacea

e Coridaceae Corispermaceae Cornaceae Corokiaceae Coronillaceae Correaceae Cor

rigiolaceaeCorsiaceae Corydalaceae Corylaceae Corynocarpaceae Coryphaceae Cost

aceae Cotyledonaceae CoulaceaeCoutariaceae Coutoubiaceae Crassulaceae Crescen

tiaceae Cressaceae Crinaceae Crocaceae CroomiaceaeCrossosomataceae Crotonacea

e Cruciferae Cryptaceae Crypteroniaceae Cryptocorynaceae CryptomeriaceaeCtenol

ophonaceae Cucurbitaceae Cunninghamiaceae Cunoniaceae Cupressaceae Curcumac

eae Curtisiaceae CuscutaceaeCyananthaceae Cyanastraceae Cynanchaceae Cyanella

ceae Cycadaceae Cyclanthaceae CyclantheraceaeCyclocheilaceae Cydoniaceae Cymo

doceaceae Cynaraceae Cynocrambaceae Cynomoriaceae Cyperaceae CyphiaceaeCyp

hocarpaceae Cypripediaceae Cyrillaceae Cyrtandraceae Cyrtanthaceae Cytinaceae

Dacrycarpaceae Dactylanthaceae Damasoniaceae Daphnaceae Daphniphyllaceae Das

ypogonaceae DatiscaceaeDaturaceae Daucaceae Davidiaceae Davidsoniaceae Decais

neaceae Deeringiaceae Degeneriaceae DelphiniaceaeDendrophthoaceae Columelliac

eae Detariaceae Dialypetalanthaceae Dianellaceae Dianthaceae DiapensiaceaeDicha

petalaceae Dichondraceae Diclidantheraceae Dicrastylidiaceae Dictamnaceae Didiere

aceae DidymelaceaeDidymocarpaceae Diegodendraceae Diervillaceae Digitalidaceae

e Dilatridaceae Dilleniaceae Dioaceae DionaeaceaeDioncophyllaceae Dioscoreaceae

Diosmaceae Diospyraceae Dipentodontaceae Diphylleiaceae Diplolaenaceae Dipsaca

ceaeDipterocarpaceae Dirachmaceae Disanthaceae Diselmaceae Dodonaeaceae Do

mbeyaceae Donatiaceae DorsteniaceaeDortmannaceae Doryanthaceae Drabaceae D

racaenaceae Dracontiaceae Drimyidaceae Droseraceae DrosophyllaceaeDryadaceae

Duabangaceae Duckeodendraceae Dulongiaceae Durantaceae Durionaceae Dysphani

aceae

Ebenaceae Eccremidaceae Ecdeiocoleaceae Echiaceae Echinopaceae Ehretiaceae Ela

eagnaceae ElaeocarpaceaeElegiaceae Elatinaceae Ellisiaceae Ellisiophyllaceae Elodea

ceae Elytranthaceae Embeliaceae Emblingiaceae EmmotaceaeEmpetraceae Encepha

lartaceae Engelhardtiaceae Enhalaceae Epacridaceae Ephedraceae Ephemeranthacea

eEpilobiaceae Epimediaceae Eragrostidaceae Eremolepidaceae Eremosynaceae Erica

ceae Erinaceae EriocaulaceaeEriogonaceae Eriospermaceae Erodiaceae Erycibaceae

Eryngiaceae Erysimaceae Erythroniaceae ErythropalaceaeErythrospermaceae Erythro

xylaceae Escalloniaceae Eschscholziaceae Eucomidaceae Eucommiaceae Eucryphiace

aeEuonymaceae Eupatoriaceae Euphorbiaceae Euphrasiaceae Euphroniaceae Eupo

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matiaceae Eupteleaceae EuryalaceaeEustrephaceae Euthemidaceae Eutocaceae Exb

ucklandiaceae Exocarpaceae

Fabaceae Fagaceae Falcatifoliaceae Ferulaceae Festucaceae Ficaceae Ficoidaceae Fit

zroyaceae FlacourtiaceaeFlagellariaceae Flindersiaceae Foetidiaceae Forestieraceae

Fothergillaceae Fouquieriaceae Fragariaceae FrancoaceaeFrangulaceae Frankeniacea

e Fraxinaceae Fraxinellaceae Freycinetiaceae Fritillariaceae Fuchsiaceae Fumariacea

eFunkiaceae

Gaiadendraceae Galacaceae Galanthaceae Galaxiaceae Galedupaceae Galeniaceae G

aliaceae Garciniaceae GardeniaceaeGardneriaceae Garryaceae Geissolomataceae Ge

itonoplesiaceae Gelsemiaceae Geniostomaceae GentianaceaeGeonomataceae Geosir

idaceae Geraniaceae Gerrardinaceae Gesneriaceae Gethyllidaceae Gilliesiaceae Gina

lloaceaeGinkgoaceae Gisekiaceae Gladiolaceae Glaucidiaceae Glechomaceae Glinace

ae Globulariaceae Gnaphaliaceae GnetaceaeGoetzeaceae Gomortegaceae Gomphiac

eae Gomphrenaceae Gonystylaceae Goodeniaceae Gordoniaceae GouaniaceaeGoup

iaceae Gramineae Gratiolaceae Grewiaceae Greyiaceae Grielaceae Griseliniaceae Gr

onoviaceae GrossulariaceaeGrubbiaceae Guaiacanaceae Guamatelaceae Guettardac

eae Gunneraceae Gustaviaceae Guttiferae GyrocarpaceaeGyrostemonaceae

Hachetteaceae Haemanthaceae Haemodoraceae Halesiaceae Hallieraceae Halocarpa

ceae Halophilaceae HalophytaceaeHaloragaceae Hamamelidaceae Hanguanaceae H

aptanthaceae Harmandaceae Hebenstretiaceae HectorellaceaeHederaceae Hedyosm

aceae Hedyotidaceae Hedysaraceae Heisteriaceae Heleniaceae Heliamphoraceae H

elianthaceaeHelianthemaceae Helichrysaceae Heliconiaceae Helicteraceae Heliotropi

aceae Helleboraceae Heloniadaceae HelosidaceaeHelwingiaceae Hemerocallidaceae

Hemimeridaceae Hemiphylacaceae Henriqueziaceae Henslowiaceae Hermanniaceae

Hernandiaceae Herniariaceae Herreriaceae Hesperocallidaceae Heterantheraceae He

teropyxidaceae HeterostylaceaeHibbertiaceae Hibiscaceae Hilleriaceae Himantandra

ceae Hippocastanaceae Hippocrateaceae HippomanaceaeHippophaeaceae Hippurida

ceae Hirtellaceae Holacanthaceae Homaliaceae Hopkinsiaceae Hoplestigmataceae H

ordeaceaeHornschuchiaceae Hortensiaceae Hortoniaceae Hostaceae Houstoniaceae

Huaceae Huerteaceae HugoniaceaeHumbertiaceae Humiriaceae Hyacinthaceae Hyd

atellaceae Hydnoraceae Hydrangeaceae Hydrastidaceae HydrillaceaeHydrocharitacea

e Hydrocotylaceae Hydrogetonaceae Hydroleaceae Hydropeltidaceae Hydrophylacac

eae HydrophyllaceaeHydrostachyaceae Hymenocardiaceae Hypecoaceae Hyperanth

eraceae Hypericaceae Hypopityaceae HypoxidaceaeHypseocharitaceae

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Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta

ceae ImperatoriaceaeInocarpaceae Inulaceae Ionidiaceae Iriarteaceae Iridaceae Irvi

ngiaceae Isnardiaceae Isophysidaceae IteaceaeStrasburgeriaceae Ixiaceae Ixioliriace

ae Ixonanthaceae

Jalapaceae Jamboliferaceae Japonoliriaceae Jasionaceae Jasminiaceae Johnsoniaceae

Joinvilleaceae JuglandaceaeJulianiaceae Juncaceae Juncaginaceae Juniperaceae Jus

siaeaceae Justiciaceae

Kadsuraceae Kaliphoraceae Kalmiaceae Kaniaceae Kiggelariaceae Kingdoniaceae Kin

giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae

Krameriaceae

Labiatae Lacandoniaceae Lachenaliaceae Lacistemataceae Lactoridaceae Lactucaceae

Lagerstroemiaceaee LamiaceaeLampsanaceae Lanariaceae Langsdorffiaceae Lantan

aceae Lapageriaceae Lardizabalaceae Lasiopetalaceae LauraceaeLawsoniaceae Laxm

anniaceae Lecythidaceae Ledaceae Ledocarpaceae Leeaceae Leguminosae Leitneriac

eae LemnaceaeLennoaceae Lentibulariaceae Lentiscaceae Leoniaceae Leonticaceae

Lepidobotryaceae LepidocarpaceaeLepidocaryaceae Lepidocerataceae Leptaulaceae

Leptospermaceae Lepturaceae Lepuropetalaceae LeuctenbergiaceaeLibocedraceae

Licaniaceae Ligustraceae Lilaeaceae Liliaceae Limeaceae Limnanthaceae Limnocharit

aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace

ae Linderniaceae Lindleyaceae Linnaeaceae LippayaceaeLiriaceae Liriodendraceae Li

ssocarpaceae Littorellaceae Loasaceae Lobeliaceaee Loganiaceae LomandraceaeLoni

ceraceae Lopeziaceae Lophiocarpaceae Lophiolaceae Lophiraceae Lophophytaceae L

ophopyxidaceae LoranthaceaeLotaceae Lowiaceae Lupulaceaee Luxemburgiaceae Lu

zuriagaceae Lychnidaceae Lyciaceae LyginiaceaeLygodisodeaceae Lysimachiaceae Ly

thraceae

Macarisiaceae Mackinlayaceae Madiaceae Maesaceae Magnoliaceae Malaceae Male

sherbiaceae MalortieaceaeMalpighiaceae Malvaceae Mangiaceae Manicariaceae Ma

paniaceae Marantaceae Marathraceae MarcgraviaceaeMartyniaceae Mastixiaceae

Matricariaceae Maundiaceae Mayacaceae Medeolaceae Medusagynaceae Peridiscac

eaeMelaleucaceae Melampyraceae Melanophyllaceae Melanthiaceae Melastomatac

eae Meliaceae Melianthaceae MelicaceaeMeliosmaceae Melittidaceae Melochiacea

e Memecylaceae Mendonciaceae Menispermaceae Menthaceae MenyanthaceaeMe

nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

47

Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

Nypaceae Nyssaceae

Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

ceae Papilionaceae Pappophoraceae PapyraceaeParacryphiaceae Parasitaxaceae Par

ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

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hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

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cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

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ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 9: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

9

Indeed there is no reason other than convention or convenience why any group

should not be segregated into several smaller monophyletic groups or merged to

produce a larger unit we can talk about one large thing or about several smaller things

Thinking about aspects of size findings in ethnobiology and cognitive psychology can be

used to suggest that a moderate number - probably fewer than 500 - of families is a

reasonable goal at which to aim and that groupings of taxa throughout any system

should be rather small in size (eg Berlin 1992 Stevens 1994 1997) Major systems such

as those of Linnaeus and Bentham and Hooker were constructed explicitly so as to ease

the burden on the memory (Stevens 1997 2002 see also Scharf 2007) the latter in

particular ensuring that all groups in their classification were relatively small often

containing three to eight immediately subordinate taxa - but by no means all their

groups were formally named Along the same lines Burtt (1977b) suggested that the

number of names at any rank should be at most one third those at the immediately

lower rank - and monotypic taxa might not need a formal name Consistent with such

ideas a fairly broad view of families and orders is taken here whenever the constraints

of monophyly and other criteria used when constructing classifications (see above)

permit The APG system is thus a convention consistent with ideas such as those

expressed by Backlund and Bremer (1998) for similar conventions see eg the Grass

Phylogeny Working Group (2001) and for fungi Hibbett et al (2007)

There are other views Thus Takhtajan (1997) has suggested that smaller families

are more natural This is incorrect Monophyletic groups that include fewer taxa -

Takhtajans smaller families - do not necessarily have more apomorphies than larger

groups even if members of smaller groups will always have more features in general in

common than the larger group that includes them That is they will have their

apomorphies their unique features as well as progressively more plesiomorphies

features found both in the small groups and in the larger clades of which they are a part

having more features in common is indeed one common meaning of more natural

(Note that the implication of the word natural has long been a group of the kind

[usually unspecified] that I think should be recognised and if other botanists had

different ideas too bad their groups were necessarily not natural [see Bather 1927] It

is thus rarely a helpful word and is not used here) Furthermore if this approach is

adopted we will find a slippery slope ahead By this kind of argument all families should

be very small since their members will have a great deal in common and so will be

most natural However as families (for example) are split the relationships that are

evident between the segregates and that were responsible for their being placed in a

single family in the first place will seem to necessitate the recognition of a new order

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

10

etc as is evident in Takhtajans own work - general taxonomic inflation is the result (see

also comparable suggestions in a cladistic context for Brassicales in particular - Ronse de

Craene amp Haston 2006) Such splitting is also questionable when teaching and learning

families since the student needs to understand the system as a whole However for

some genera removed from the families that until now have included them the

phenetic-classificatory-phylogenetic structure in their new home may mandate the

recognition of small families On the other hand Takhtajans suggestion that narrowly

defined families are more useful for phylogenetic studies may be true Indeed I have

more than once regretted prematurely combining groups whether species (in the

context of monographic work) or families (in the course of preparing these notes) but

this is largely a separate issue

Van Steenis (1978) Philipson (1987b) and others have rightly questioned the value

of splitting a group when ideas of the relationships of its constituent members have not

changed - that is very good reasons have to be provided for splitting a family if the

genera within it remain part of the same clade rather than belonging to another clade

Thus APG (2003) broadened the circumscription of Malvaceae because of the

parapolyphyly of some of the families that had historically been associated with it (Judd

amp Manchester 1997 Alverson et al 1999 Bayer et al 1999) These families particularly

Tiliaceae and Sterculiaceae were not at all easy to distinguish their close relationship

(see eg Brown 1814) and overall similarity had long been conceded and to some

workers at least their combination has come as something of a relief Although most of

the larger clades within Malvaceae sl remain difficult to distinguish even with flowers

Cheek (2007) opts for a wholesale and novel dismemberment into ten families

however the very good reasons for doing this are wanting

The same principles are of course applicable when it comes to dividing genera little

other than a headache is gained by splitting genera such

as Drosera and Gnetum (Doweld 2000) as has recently been proposed Thus if an

established genus divides into two (or more) clades this is not a signal for recognising

two groups at the same level - so here it could be argued that the dismemberment

of Pterostylis (Jones amp Clements 2002b) was somewhat unfortunate Along the same

lines if a newly-discovered taxon is sister to an existing named taxon - say a genus - this

does not necessitate the description of a separate genus for the newly described species

(cf Davis 2002) (Of course some [eg Thorne 1976] have suggested that the sizes of

gaps between groups at the same rank should be similar but any principle like this is

inherently flawed since morphological gaps are more unstable than phylogenetic

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

11

relationships and applying it across all flowering plants would both be difficult and cause

substantial changes - see also below) Humphreys and Linder (2009) provide a well-

documented survey of generic concepts in plants which the reader should consult they

note that generic limits (broad versus narrow) have oscillated historically and that

currently larger genera tend to be recognised because studies tend to be on a broader

scale than in the past

Note that invoking similarity or difference - whether qualified (considerable

similarities substantial differences) or not - in a cladistic context as justification for

combining or splitting taxa is not a particularly strong argument (see eg

CardiopteridaceaeStemonuraceae - Karingrehed 2002c) Similarity and difference can

neither be defined precisely since what may seem to be substantial similarities to me

may not to the next person nor are they likely to be stable in the face of our changing

knowledge of morphology and what might be synapomorphies

I might have prefered to merge some families recognised here or split others but

by and large I do not think my own preferences matter very much - and I take the same

position with regards to comparable preferences expressed by others Indeed the

bottom line is that in flagged hierarchies of the kind used here the limits of any

monophyletic unit generally taught and discussed particularly other than species can

be established only by convention and consensus (eg Stevens 2002 2006a Entwisle amp

Weston 2005) This is rather different from the reasonable observation that the

phylogeny itself cannot be achieved by consensus (Thorne 1976) for in most cases there

is a fact of the matter when it comes to relationships Given the increasing support for

the outlines of angiosperm phylogeny a stable consensus classification based on this

phylogeny seems attainable Indeed in addition to providing current ideas of

relationships of seed plants in a synthesised form this site is part of an attempt to build

such a consensus about the circumscription of taxa (see APG 1999 2003 2009 Grass

Phylogeny Working Group 2001 Mabberley 2008) Reaching such a consensus is vital

since what we know of angiosperm phylogeny allows a very large number of

classifications to be based on it and as we find out more the number of possible

classifications increases greatly Unfortunately however nature does not dictate what

the classification should be All classifications are constructed by humans to

communicate particular aspects of groups and relationships Our goals as systematists

are surely to produce robust hypotheses of relationships to understand the evolution of

morphology and the like - but not to argue ad nauseam whether something should be

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

12

a family or a subfamily That way surely lies madness and worse the discredit of our

discipline

There are similar issues whatever naming system is used Thus in phylogenetic

naming (Baum et al 1998 for an example but cf Baum et al 2004 for the PhyloCode

see Cantino amp De Queiroz 2006) an unflagged hierarchy is used in which any

terminations of names used are uninformative about the relative position of taxa If one

adopts the principle of phylogenetic naming one indeed does not have to worry about

the nomenclatural consequences caused by lumping or splitting any well-supported

clade can be named without affecting the name of more or less inclusive clades

Unfortunately unflagged hierarchies have very serious deficiences as communication

devices because they lack one aspect essential in language biological or otherwise -

they contain no intrinsic information about the relationships of the group in question to

others (eg Pfeil amp Crisp 2005 Stevens 2006a) Recent suggestions for using prefixes like

Apo- and Pan- to PhyloCode names will however allow limited information of this

kind to be conveyed but only as it pertains to individual branches and current

proposals do not even mandate that the prefixes be employed consistently In any

event such proposals simply prevent the potential tripling of the number of quite

different names used to describe different aspects of a phylogenetic tree over those

used to name monophyletic groups pure and simple In general where n is the number

of extant species in a group the number of clades in such a group = n-1 (Species will

also need names too for their names see Dayrat et al 2008) Importantly here too

consensus over the clade names commonly learned by students and used in herbaria is

needed otherwise communication will be impeded the names themselves will provide

no guidelines as to which should be chosen The situation is of course more complicated

than this Terminations that convey ideas of rank in a phylogenetic classification can also

be used in phylocode names - however there they will carry no implications of rank

How they will be used is another matter of course

Of course there are other philosophies of classification and some still prefer

evolutionary classifications There classificatory principles differ substantially from those

followed here eg the recognition of paraphyletic taxa may be permitted however

detailed reasons for prefering the taxa that are recognised are rarely given although

nature and natural groups are often mentioned (cf Stuessy amp Koumlnig 2008) For

summaries of commonly used systems see Brummitt (1992) and Mabberley (2008)

new evolutionary systems appear every year or so However even those who allow or

promote the recognition of paraphyletic groups (eg Grant 2003 Thorne 2007

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

13

Heywood et al 2007) may find it of interest to examine a system recognizing only

monophyletic groups it provides a rather different understanding of evolution

Evolutionary classifications in general try and combine phylogeny and

morphological gaps although that is no easy thing to do - it is akin to combining chalk

and cheese (for an attempt to make this impossible task seem to be more objective see

Stuessy amp Koumlnig 2008) However there is a principle from evolutionary classification

that is relevant and I think quite useful here The size of the gap between two groups

tends to be inversely proportional to the sizes of the groups involved (Davis amp Heywood

1963) One can imagine a situation in which a large group is formally divided even

although the distinguishing characters of the two are weak whereas a smaller group

similarly divisable will be left intact

To summarize If hypotheses of phylogeny remain stable we should be able to base

a stable classification on that phylogeny and then get on with our work that is testing

the phylogenies we have elucidating phylogenies in areas where relationships are

unclear studying the evolution of morphology describing species etc In this context

the spread of the Angiosperm Phylogeny Group system (see below) and its widespread

utilisation in technical literature also floras (eg van der Meijden 2005) textbooks (eg

Simpson 2006 Judd et al 2007 [third edition]) dictionaries (Mabberley 2008) more

popular literature (eg Souza amp Lorenzi 2005 Spears 2006) and as an outline for a new

herbarium sequence (Haston et al 2007) is gratifying The posibility that one might be

able to develop a stable phylogeny-based classification of families and in particular

orders represents a dramatic turn-around from the pessimistic attitude about such

higher-level groupings expressed by Davis and Heywood (1963) and Thorne (1976) the

latter even suggesting that we should bury forever the metaphor of the phylogenetic

tree as highly unrealistic (ibid p 56) Returning to Godfray and Knapps (2004) users

of classifications who want a stable informative and accessible classification that

enables easy identification - unfortunately they want cake with everything and cannot

get it - these pages attempt to satisfy as many of their needs as possible but phylogeny

and monophyly are the primary shapers of the APG classification

On this classification in particular

Here I very largely follow the Angiosperm Phylogeny Group classification (APG

2003) Any differences are not to be interpreted as differences in principle simply that

new phylogenies continue to be published and here I attempt to provide an overview

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

14

of current ideas of higher-level relationships of seed plants The Angiosperm Phylogeny

Group classification is based on well-supported relationships evident in the numerous

molecular studies that began to appear in the late 1980s (see APG 1999 for the

principles underlying the classification) but recent phylogenetic work does not

contradict the major outlines of the trees used by APG II (2003) or even those in APG

I (APG 1999) The APG classification has for the most part been conservative and I

am similarly conservative here For the names and authorities I follow APG III (2009)

although the names of the authors may not always be correct these being in something

of a state of flux for the superordinal names used see Chase and Reveal (2009)

although classification at this level here is not exhaustive - only clades with several

orders have superordinal names

Some changes in our ideas of relationships and hence in the clades we talk about

are particularly likely in parts of Lamiales Caryophyllales and Malpighiales for example

although this will not affect the orders themselves Thus some changes to clade

circumscriptions are to be expected although I do not expect them to be substantial

but changes are neither a defect of cladistics nor a necessary consequence of the use of

molecular data Clades are hypotheses of relationships and as hypotheses they may be

overturned However work since APG I has not suggested other than minor changes

in the compositions of the orders even if the odd genus or even family is turning out to

be seriously misplaced - recent examples are Hydatellaceae (from monocots-Poales to

Nymphaeales Saarela et al 2007) Guamatelaceae (from Rosales-Rosaceae to

Crossosomatales Oh amp Potter 2006) and Perrottetia and Bhesa (from Celastrales-

Celastraceae to Huerteales and Malpighiales respectively Zhang amp Simmons 2006) The

main changes have been clarification of the relationhips of individual families or groups

of families that were of uncertain position eg of Chloranthaceae (Moore et al 2007)

Ceratophyllaceae (Jansen et al 2007) and unplaced asterid II families (Winkworth et al

2008a) As mentioned above phylogenies in the area of main interest in these pages are

overwhelmingly tree-like Although there is evidence for acquisition of host plant genes

by parasites (see below) even wider but inexplicable transfer of mitochochondrial

genes in plants like Amborella (Bergthorsson et al 2004 cf Goremykin et al 2009a)

and even transfers of nuclear genes (Vallenback et al 2008) these are the exception

rather than the rule

In cases where the Angiosperm Phylogeny Group II allowed alternatives as to the

limits of families - Papaveraceae in the broad sense or Papaveraceae plus

Pteridophyllaceae plus Fumariaceae Proteaceae in the broad sense or Proteaceae plus

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

15

Platanaceae - the choices made here follow common usage eg as in textbooks like

Judd et al (2007) and Simpson (2006) and particularly in the new edition of

Mabberleys The Plant Book (Mabberley 2008) This latter is itself an attempt to reflect a

consensus the result of taking the opinions of botanists at several meetings A largely

similar consensus is reflected in the APG III classification (APG 2009) where you will

see that alternative classifications have been dispensed with For many people the

existence of such alternative classifications will simply confuse so agreement over

which groupings to use when alternatives are permitted and then dispensing with the

alternatives that are not commonly used seems reasonable I am making the few

changes that are needed to bring the classification here in line with these others but

only gradually since there is not enough time in each day

As already mentioned higher-level relationships in general and the composition of

orders in particular have always presented something of a challenge to systematists

(eg Davis amp Heywood 1963) That being said the composition of some clades like

Apiales Crossosomatales and Pandanales is decidedly unexpected However it is

interesting to see that these higher level clades are generally accepted even in works

with different classificatory philosophies (eg Heywood et al 2007) For clades like

Malpighiales many of the family groupings within Asparagales etc attempts to find

distinctive characters have largely failed (but see Endress amp Matthews 2006a also

the Apomorphies page here) Interestingly as with families some groupings suggested

by molecular studies are supported by morphological andor chemical characters that

have long been known sometimes for over a hundred years the relationship between

Pittosporaceae and ApiaceaeAraliaceae are a case in point (Hegnauer 1969b and

references) As our knowledge of morphology and chemistry improves we can hope for

improvements in the characterisations of clades at all levels

In many cases the new family limits of the Angiosperm Phylogeny Group (see

APG 1999 2003) are not really controversial although changes from the limits

commonly accepted only a decade ago are sometimes quite dramatic (eg Wagenitz

1997) Thus the split of the old Saxifragaceae sl is necessitated by its extreme

polyphyly as also with Icacinaceaesl and Cornaceae sl However the limits and

relationships of such groups had long been considered to be unsatisfactory but until

recently there had been no real reason to prefer one proposed arrangement over

another It is generally accepted that the limits of Lamiaceae and Verbenaceae have to

be redrawn although there is as yet no compelling evidence that the redrawn taxa are

not sister taxa (there is no evidence that they are) But whatever their relationships the

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

16

content of the clades has changed considerably and incidentally they are now easier to

identify than before the decision to recognise two families is not difficult The same is

true for Salicaceae and Achariaceae (Malpighiales) two previously small families that

have received the bulk of the old Flacourtiaceae Clade and hence taxon limits remain

difficult around Theaceae (Ericalessee alsoSladeniaceae Pentaphylacaceae)

and Euphorbiaceae (Malpighiales see

also Phyllanthaceae Picrodendraceae Putranjivaceae etc) Here current groupings

may not represent quite such dramatic changes in our understanding of relationships

Indeed parts of the old Euphorbiaceae that were separated may yet go back together

particularly Phyllanthaceae and Picrodendraceae (see Wurdack et al 2004 Davis et al

2005) even if Theaceae should indeed be dismembered (Geuten et al 2004) However

given that there is no molecular evidence that currently warrants combining all the

segregates of Euphorbiaceae and even if some do come together the clades the

families represent suggest novel groupings not recognised in current classifications

Furthermore Rafflesiaceae appear to be embedded within Euphorbiaceae s str (Davis

et al 2007) so the family is divided Of course maintaining even a moderately narrowly-

circumscribed Euphorbiaceae would entail reducing the iconic Rafflesiaceae to

synonymy Relationships in core Caryophyllales especially

around Phytolaccaceae andNyctaginaceae and also Portulacaceae are incompletely

understood and refashioning of taxon limits will doubtless be needed as cladistic

relationships become apparent (eg see Nyffeler 2007 Ogburn 2007a b) Some

groupings in the old Icacinaceaeand Olacaceae also remain unclear

The discovery of the relationships of parasitic and aquatic groups have presented a

particular challenge to systematists Morphologically some of these plants are so highly

modified that interpretation of the plant body in conventional terms is difficult or even

impossible Thus parasitic groups (broad-leaved angiosperms only) such as Rafflesiaceae

are hard to place since both the vegetative body and the flowers are changed almost

beyond recognition (flowers of rafflesiaceae are a good example of gigantism - Davis et

al 2007) Furthermore plastid gene sequences may be difficult or impossible to obtain

the chloroplast DNA in particular being highly degraded and the rate of molecular

change in general may be high so the problem of long-branch attraction is serious (eg

Duff amp Nickrent 1997 Nickrent et al 1998 Caddick et al 2002a G Petersen et al

2006b) Echlorophyllous sapromycoheterotrophic taxa mostly known from monocots

present similar problems However progress is being made placements for

Apodanthaceae Rafflesiaceae Mitrastemonaceae Cytinaceae and Cynomoriaceae have

recently been suggested (Barkman et al 2004 2007 Davis amp Wurdack 2004 Nickrent et

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

17

al 2004 Davis et al 2007) it seems likely that Burmanniaceae sl are polyphyletic but

both parts are in Dioscoreales (Merckx et al 2006) and relationships within the largely

hemiparasitic Santalales are also gradually being clarified (Maleacutecot 2002) The inclusion

of parasitic taxa in general molecular analyses can cause conniptions (eg Nickrent et al

2004 Davis et al 2004 Chase et al 2006 G Petersen et al 2006b) and there can be

horizontal transmission of genes (eg Davis amp Wurdack 2005 Vitaceae to Rafflesiaceae

Barkman et al 2007 the mitochondrial atp1 gene commonly moves)

Similarly in water plants neither vessels in particular nor much xylem in general is

needed leaves are highly modified and water-mediated pollination if adopted may

well be associated with major changes in floral morphology Here too recent molecular

studies suggest that aquatic groups with hitherto problematic relationships may find

homes Thus Podostemaceae are close to Clusiaceae and Calophyllaceae and are sister

to Hypericaceae (Malpighiales Kita amp Kato 2001) Hydatellaceae which used to be in

Poales are part of Nymphaeales (Saarela et al 2007) and Hydrostachyaceae may be

close to Hydrangeaceae (Cornales Xiang et al 2002) or perhaps should be placed in

Lamiales note that in the first two cases in particular there are morphological and

chemical features that support such a move If Podostemaceae are indeed close to

Clusiaceae I look forward to seeing hypotheses to explain how the dramatic changes in

the vegetative body that have made Podostemaceae so problematic for generations of

systematists took place That conventional wisdom has trouble in understanding or

explaining how the morphologies of groups like Clusiaceae and Podostemaceae can be

related is largely a problem with conventional wisdom Ceratophyllaceae seem to be

finding a position as sister to eudicots (see Moore et al 2007) but their morphology is

so derived that there is no morphological evidence of which I am aware for this

relationship There are also similar although less extreme morphological problems with

many plants pollinated by wind

In such situations especially with some water plants mycoheterotrophs and

parasites a variety of characters may be affected and particularly under such

circumstances groups have tended to be formed using a few or even only a single

character that seems to provide evidence of relationships Add to this the tendency to

weight some characters particularly strongly on a priori grounds and the result was

either the recognition of conglomerate taxa such as Amentiferae which now appear to

be highly polyphyletic or the segregation of families like Plantaginaceae s str (now

much expanded) and Leitneriaceae (now in Simaroubaceae) - all of these are more or

less wind pollinated

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

18

Even although relationships of many parasitic and aquatic groups are being

clarified and at least some of the proposed phylogenetic positions seem to be well

supported it is indeed often still very difficult to understand how such groups have

evolved from their more morphologically conventional relatives However we have had

all sorts of preconceptions as to how characters should evolve Thus we used to

assume that features like highly scalariform vessel perforation plates or the complete

absence of vessels or a flower with an androecium that had many stamens a superior

ovary or separate petals were necessarily plesiomorphic or primitive and conversely

a simple perforation plate and the presence of vessels an androecium with few

stamens inferior ovary or petals that were connnate were almost necessarily

apomorphic or advanced (but cf eg Stebbins 1951) Such assumptions are incorrect

(eg Soltis et al 2005b) Carpels may become secondarily free carpels may fail to close

the seeds then developing outside the confines of the carpel as in some Aspagaraceae-

Nolinoideae Violaceae Berberidaceae Malvaceae-Sterculioideae etc) in Peliosanthes

teta perhaps the only species in Peliosanthes(Asparagaceae-Nolinoideae) the ovary

varies from superior to inferior (Jessop 1976 species limits here need close

investigation see also Kuzoff et al 2001 and Soltis amp Hufford 2002 Saxifragaceae

Apiales etc) many-seeded carpels can evolve from few-seeded carpels

(Razafimandimbison et al 2008) monoecy may be derived from dioecy (Schaefer amp

Renner 2010 and references) Classic studies such as those by Babcock (eg 1947)

on Crepis that assumed that evolution - in this case of the karyotype in particular - was

unidirectional have needed comprehensive re-evaluation (Enke amp Gemeinholzer 2008)

Most if not all characters have reversed andor evolved in parallel as is clear in the

discussion of ovary position here especially in Poales Saxifragales and Asparagales and

also in comments on such morphologically distinctive taxa (when compared with their

immediate relatives) such as Menyanthaceae and especially Pittosporaceae Parallel

evolution may occur even at the level of amino acid substitution as in the independent

acquisition of the phosphoenolpyruvate carboxylase (pepC) gene in C4 photosynthesis in

grasses (Christin et al 2007b see also Blaumlsing et al 2000)

Thinking of how such characters have been used in the past in classifications at the

generic level the results have been similar to those just discussed above for parasitic

plants and others Relying too much on animal pollination syndromes to mark generic

boundaries has all too often led to taxa that are highly unsatisfactory phylogenetically

(see eg Acanthaceae Bignoniaceae Campanulaceae Ericaceae Lamiaceae

Orchidaceae) and over-reliance on characters of fruit and seed (see eg Brassicaceae

and Apiaceae) has also led to unsatisfactory generic limits Again the more general

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

19

problem is the use of one or a very few characters that have been weighted a priori to

structure classifications (see also Garciacutea et al 2009) although it is perhaps not

surprising that such an approach sometimes results in the delineation of taxa that have

indeed turned out to be quite distinct (cf eg Burtt Astragalus versus Oxytropis)

Taxa recognized here are monophyletic yet many are polythetic at the

morphological level that is they lack unique features characterizing (defining) all and

only members of that family they can be recognised phenetically only by the unique

combinations of characters that they posess This is the result of evolution any

synapomorphy characterizing a taxon may be lost or modified beyond easy recognition

in some of its members or the synapomorphy may appear to be identical to a feature

that has evolved in parallel in a quite unrelated plant That plant groups are polythetic is

almost as much a feature of monophyletic taxa as the paraphyletic taxa common in

evolutionary classifications Some families now include substantial variation as

phenetically distinct derived groups are placed in their proper phylogenetic position -

examples are Ericaceae which include the erstwhile Empetraceae a wind-pollinated

group that is florally very different and the overwhelmingly large-flowered and animal-

pollinated Plantaginaceae which now include several derived small-flowered aquatic

and wind-pollinated groups that were previously placed in separate families

Thus some families as delimited here may not be easy to recognize However

remember that detecting relationships - use whatever characters you can even if they

are not obvious - and naming a plant - focus on easy-to-see characters that may not

reflect relationships - are quite different problems Taxa although natural may not be

readily recognizable indeed it was in exactly this context that Lamarck worked out the

basic principles of writing dichotomous keys in 1778 (see Scharf 2007 for keys and the

like) Of course Lamarcks idea of nature was very different from ours - he thought that

there was some kind of continuum of form on which living organisms were to be

situated with no real gaps anywhere - but this meant that his genera (for example)

might well not be sharply distinct from each other even if each were part of the real

continuum that was life Lamarcks separation of the establishment of relationships and

groupings in a system and identification of the plants in that system have been widely

accepted (but cf eg Godfray amp Knapp 2004 many of the contributors to Heywood et

al 2007) Perhaps the best way of identifying plants at the family level is by well-made

multiple access keys as in Watson and Dallwitz (1992a onwards family limits there may

differ substantially from those adopted here) Multiple access keys free users from the

constraints of dichotomous keys in which particular characters are needed at each step

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

20

of the identification process before there can be further progress Instead those

characters that happen to be evident on a specimen can be used in whatever order is

convenient when linked to illustrations glossaries etc their power is enormous (see

Dallwitz et al 2000 [2006] for the principles underlying their construction and use)

Nevertheless dichotomous keys such as those of Hutchinson (1973) and Franz Thonner

(Geesink et al 1981) have their uses Of course taxa coming out adjacent in keys may

well not be at all related

(When identifying large numbers of plants even more efficient than either style of

identification and certainly lots more fun is sight identification Unless you have a

photographic memory you have to build up your knowledge of comparative plant

morphology - on which the ability to make accurate identifications depends - by

repeated observation When faced with an unknown plant I always look for leaf teeth

and stipules distinctive hair types especially stellate and T-shaped hairs glands and

punctations of any sort the presence of latex or other exudate and check leaf insertion

smelling crushed leaves can also be helpful In this context nodal anatomy can usually

be checked using a razor and a hand lens or even simply carefully examining leaf scars

The short paragraphs added after most families may help in confirming familial

identifications)

For the record and for the little that it is worth there are 4 orders and 13 families

of gymnosperms characterised on these pages and together they include some 82

genera and 947 species For angiosperms comparable figures are 56 orders 445

families 13208 genera and 261750 species (of which monocots include 11 orders 89

families 2759 genera and 52760 species) Note however that higher mathematics

was never my strong point and anyway these are pretty meaningless figures even for

species which many (but not all) might concede smacked slightly more of reality than

other taxa estimates range as high as 422000 (Govaerts 2001) Furthermore numbers

of genera and species change daily Nevertheless as emphasized here families are

useful in teaching we as a community can ensure that their limits remain largely stable

and by concentrating on relatively few of them one can gain some familiarity with much

of the worlds flora For summaries of the sizes of orders and families and of the general

arrangement followed in these pages see also the Statistics page itself

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

21

A PHYLOGENETIC TREE OF SEED PLANTS (THE MAIN TREE)

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

22

The Main Tree above is a conservative summary of well supported relationships

between the major clades of extant seed plants Reading - interpreting and

understanding - such a phylogenetic tree is not necessarily easy so reading a tree may

require a bit of practice OHara (eg 1988 1992) provides an accessible introduction to

thinking and talking about history in the context of looking at phylogenetic trees More

recently Krell and Cranston (2004) Crisp and Cook (2005) and others have emphasized

how careful one must be when interpreting and talking about evolution in ladderized

trees in particular and phylogenetic trees in general Gregory (2008) provides a

particularly detailed discussion of this problem In ladderized trees the smaller (in terms

of numbers of terminals) sister taxon is consistently shown on the same side at every

node thus the trees tend to be pectinate like the teeth of a comb however the

horizontal axis that is as a result so evident has no polarity or particular direction The

use of the adjective basal when discussing phylogenetic trees is especially dangerous

(see also Wojciechowski et al 2004 D Soltis et al 2005b) When I use the term and the

context is not otherwise clear I am referring to the pectinations at the base of a

ladderised tree Remember that when talking about sister taxa one can never be basal

to the other or older or younger although members of one can be more derived - in the

sense of having more apomorphies - than the other (but even then one has to be

careful) The words primitive and advanced and lower and higher should

practically never be used especially when talking about taxa Amborellaceae and

Pinaceae are sister to all other flowering plants and Pinales and are basal to the crown

groups of all other flowering plantsPinales respectively but that does not mean they

are thereby primitive Pinaceae in particular have numerous apomorphies The word

plesiomorphic is far less loaded than primitive and can be used to talk about

individual characters

Most trees here have been more or less ladderized as is the one above The

ladderization may be imperfect for example you can see that on the Main Tree

asterids with ten orders follow rosids and relatives which have seventeen although

this is in part because of the currently unresolved nature of relationships between the

main clades of core eudicots (this is likely to change soon) The Main Tree could have

been drawn with Amborella Acorus or a host of other taxa at the far right without

offending any relationships phylogenetic trees are like mobiles the only fixed points

being the nodes Despite the problems just mentioned pectination interpreted

carefully has its value and it is not an arbitrary process As one reads the terminals of a

pectinate tree from left to right adjacent terminals will be separated by apomorphies

that will cumulate in an appropriate fashion When thinking of a book or a herbarium

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

23

sequence (see Haston et al 2007) this is of some value Nymphaeales and

Austrobaileyales are here adjacent on the tree but they could be separated by

hundreds of families in the sequence merely by rotating only the node from which

Nymphaeales and their sister group (all other angiosperms minus about 75 species)

arise If adjacent in a book or herbarium or as in these summary pages then it is

relatively easy to relate their characters both apomorphic and plesiomorphic but if

separated by hundreds of pages or two floors in a large building then it is less easy to

get anything from the sequence (Since all orders in the pages of the main part of this

site are preceded by the apomorphies of all nodes immediately below them in the seed

plant phylogeny and because of the linkages that have been built in to the site this

problem is much less serious there there is no sequence) So the particular sequence of

all seed plants that is based on a phylogenetic classification but is to be used in herbaria

or any other place where taxa have to be arranged linearly as in this part of APweb

will be one that maximizes the number of taxa that are both successive branches of the

tree and placed successively in the sequence Since specimens are generally filed under

families the outline of a new family sequence for arranging herbaria and books can now

be suggested (Haston et al 2007) and such a standardized sequence will further help

teaching and learning about plants

ON THINKING ABOUT APOMORPHIES

Identifying apomorphies is important because understanding the evolution of

morphology in the broad sense - ie including anatomy chemistry etc - in the context

of diversification is one of our major goals For identifying apomorphies several

preconditions must be met One needs to have an accurate well supported phylogeny

one has to have examined the right taxa both from the point of view of morphology and

molecules one has to have coded the characters right (ie delimited states

appropriately) and one has to use an appropriate model of evolution when fitting the

variation to the tree (see eg Omland 1999 Stevens 2006b) If the distributions of

apomorphies below are compared with those in other studies (eg Turgeon et al 2001

Bremer et al 2001 Endress 2001 Albach et al 2001a Judd amp Olmstead 2004 D Soltis

et al 2005b Zhang et al 2006 Judd et al 2007) the reader may find a number of

differences Although I have integrated such studies as far as is possible there are five

reasons that link to the preconditions just mentioned why there may be differences

where features are placed on trees

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

24

1 I may not have found all the information about a particular character there may be

disagreement over its interpretation or I have added information

2 The sampling of nearly all molecular studies is very incomplete (see Salisbury amp Kim

2001 for problems caused by sampling) indeed it is commonplace to decry the

incompleteness of molecular sampling But not only is the sampling in molecular studies

often less than we might wish that of the morphological and chemical characters whose

evolution we are interested in understanding is also often very poor So for many

anatomical chemical and embryological characters that are confidently said to

characterise families and other groups we all too often have no idea if those characters

are applicable to the whole clade or just to a subgroup within that clade However as

our sampling improves we can locate changes on the tree more precisely if in

unexpected positions Thus is is sometimes suggested that Ericaceae have ellagic acid

(Soltis et al 2005b) in fact the little evidence we have indicates that only a small clade

of some 80 species has ellagic acid and the rest of the family - itself alone about one

third of all Ericales - does not Similarly Ericaceae are noted for their

ectendomycorrhizal associations but Enkianthus sister to all other Ericaceae

apparently lacks these (Abe 2005) Thus these mycorrhizae and their associated

distinctive hair roots are unlikely to be apomorphies of Ericaceae - however it appears

that other families near Ericaceae may also have ectendomycorrhizae (see Asai 1934)

3 I am fitting characters to a very conservative tree with many polytomies although the

nodes that are utilised are for the most part strongly supported polytomies make the

optimisation of characters that is the assigment of character state change to a

particular node on the tree notably difficult (eg Madison amp Madison 2002) In nearly

all studies of the evolution of characters distributions of characters are optimised on a

more or less fully resolved tree and the construction of supertrees may yield yet more

detailed hypotheses of relationships (for literature on supertrees see Cotton amp

Wilkinson 2007 2008) Of course some nodes on such fully resolved trees andor

supertrees may have little support and optimisations of characters on such trees may

carry correspondingly little conviction Even parts of some of the trees used here have

poor support eg relationships within aquatic Alismatales etc although I indicate

when this is the case

4 Exactly how one goes about optimising a character on a tree is critically important

Even using simple parsimony optimisations (ACCTRAN or DELTRAN ACCelerated

TRANsitions or DELayed TRANsitions) the position of synapomorphies on trees - and

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

25

hence our ideas of evolution - may differ and this is still more true if one uses maximum

likelihood or Bayesian methods and makes apparently reasonable suggestions about

weighting gains over losses (eg Donoghue amp Ackerley 1996 Cunningham et al 1998

Omland 1997 1999 Ree amp Donoghue 1999 Polly 2001 Webster amp Purvis 2001

Ronquist 2004 Crisp amp Cook 2005) Sannier et al (2007) show how in in Arecaceae that

where on a phylogenetic tree one might peg changes in microsporogenesis will depend

on the methods one uses to do this and Pedersen et al (2007) discuss the sometimes

very substantial effect of node support on the posterior probabilities of ancestral

character states Here I use parsimony optimization not always as explicit as it might be

but I have often indicated where there are particularly important uncertainties as to the

positions of particular character changes on the tree

5 Finally although I have paid quite a lot of attention to the delimitation of the

character states that make up all the characterizations I have not spent enough time on

this critical operation If we are interested in understanding evolution then fitting the

basic variation - not character states - to a tree in principle allows greater flexibility in

understanding morphology in the context of local phylogenies (see also Stevens 2000

Endress 2005c) However many character states used here are delimited globally that

is they are circumscribed in the context of the variation shown by individual characters

across all angiosperms andor in the context of classic ideas of character evolution

Character states often have arbitrary limits and serve best to communicate

information whether they are in fact suitable for either phylogenetic analysis or

understanding evolution are separate issues Studies have rather unsurprisingly

perhaps but importantly shown that dividing the one character into different sets of

states may yield differing ideas of evolution of that character (eg Lamb Frye amp Kron

2003 Hibbett 2004) When looking at trees on which character states are optimised

one should bear in mind the problems surrounding the delimitation of states (eg

Stevens 2000 2006b) and the danger of using pollen or other types - constructs

based on many characters that vary independently but which effectively get lost in

these types thus Blackmore et al (2009) decomposed the pollen types in Asteraceae

into 52 characters

I use Remanes three main criteria of homology or better similarity when

determining the basic similarity of structures on different organisms (see Remane 1952)

These criteria are special properties position and intermediates Special properties

include anatomical or chemical characters gene expression data etc Position refers

to the position of an organ with respect to landmarks on the plant Although plants are

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

26

plants and landmarks are not as fixed as one might like trying to work out the relative

positions of parts is a good way of understanding morphology so long as one does not

try to out-do Procrustes Intermediates include intermediates found on the same

plant or on different plants Intermediates may observed during development or be

adult structures Thus one may find changes during development which suggest what

the nature of a particular structure is and very different stuctures on different plants

may turn out to be similar early in development Or one can simply compare different

taxa and see that structures that appear to be very different when comparing two

different taxa can be linked morphologically by looking at other taxa

Nevertheless the use of these criteria may not yield an unambiguous answer as to

what a structure is even given a solid phylogeny and an improved understanding of

development (see Jaramillo amp Kramer 2007 for a useful discussion) As Endress (2005c)

observed a number of features - position function development shape anatomy

histology gene activity and relationships to other taxa that clearly have petals - can be

used to distinguish a petal (for example) from other floral structures if a petal does not

have one of these features is it thereby not a petal Thus Maturen et al (2005) recently

found that floral organ diversity genes (B and C) were expressed in the large white

inflorescence bracts of Cornus (see also Costa et al 2005) Peney et al (2005) noted that

not all monosulcate pollen grains in monocots have the same developmental pathway

and that as a result such pollen might not have the same ancestral state Reeves and

Olmstead (2003) suggested that the genetic mechanisms causing monosymmetry in

Lamiales and Solanales were different and Serna and Martin (2006) described similar

problems with the development of hairs in Arabidopsis when compared with that of

hairs in Antirrhinum and Solanaceae Indeed as one perhaps might expect delimitation

of states and characters does not necessarily become easier with increasing knowledge

of development etc Thus Buzgo et al (2004) Matthews and Endress (2005) and others

have shown how hard it can be to distinguish between eg prophylls and other floral

structures as their behaviour is studied during the course of the development of the

flower Similarly at what concentration is a particular secondary metabolite deemed to

be present (Waterman (2007)

The validity of the approach used here that of fitting morphological variation to a

largely molecular-based tree may be questioned However I think it rather unlikely that

well-supported molecular branches will be overturned by morphological data Indeed

analyses of morphological data alone do provide support for many of the clades evident

in molecule-only analyses and analyses with morphological and molecular data

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

27

together may lead to increased support for clades (eg Hufford 1992 Nandi et al 1998

[but adding morphological data reduces support for a number of critical clades too]

Doyle amp Endress 2000 however in none of these papers is the use of morphology

without ambiguity) It is unfortunately clear that the use of morphology alone may not

suggest problems in the phylogenetic placement of taxa that later turn out to have been

wrongly included (eg Zhang et al 1992) Thus I have been wary of putting much weight

on clades that have only morphological support but note that I have treated molecule-

based clades with low bootstrap or jacknife support values (esp below 70) or low

posterior probabilies (below 095) likewise Although I may have been mistaken in

placing so much emphasis on molecular data in terms of providing the basic

phylogenetic framework for angiosperms morphological and molecular data are only

very rarely in irreconcilably strong conflict There are indeed a few places where the

conflict seems extreme These include the relative positions of the Monimiaceae and

Hernandiaceae (Laurales) the position of Hanguanaceae (Commelinales [as here] or

Zingiberales) and of Triplostegia (is it in Dipsacaceae or Valerianaceae - see

Dipsacales) Fossils are unlikely to affect the topologies of the trees presented here but

see below for their importance in understanding morphological evolution in general and

the evolution of angiosperms in particular Nevertheless some largely reject the idea

that trees based on molecular data alone can recover phylogenetic relationships

especially when branching points are old and prefer to used trees based on analysis of

morphological data including those taken from fossils (Hilton amp Bateman 2006 Farjon

2007)

All in all however the extent of the congruence between morphological and

molecular data is impressive and heartening and many clades can be characterised

morphologically It seemed in 1998 that there were no unambiguous morphological

synapomorphies for angiosperm orders (K Bremer 2000) and this is still true if by

unambiguous is meant non-homoplasious However many orders and other clades

have synapomorphies even if these may be indistinguishable at least at the current

level of morphological and developmental knowledge from parallel occurences

elsewhere As our morphological knowledge increases so too does the number of

apomorphies

Endress and Matthews (2006a) emphasize the importance of tendencies and

developmental constraints when thinking about characters of clades within the rosids -

many characters or character combinations occur in a rather sporadic fashion within

the clade and apparently notably less frequently outside it There are a number of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

28

examples of tendencies here and in the individual order pages such as the features

enclosed by parentheses in groups above the level of family eg see (cuticular waxes

as aggregated rodlets) for the commelinids and the discussion of the distribution of

polyandry that is flowers with numerous stamens in the asterid I + II groups etc

Tendencies may involve more than single characters When thinking of the

associations of features like integument thickness and vascularization of the integument

(Endress amp Matthews 2006a) one can imagine fairly simple developmental

preconditions being operative As Endress and Matthews (2006a) note it is difficult to

imagine vascular tissue developing in an integument only two cells thick Changes here

would be loosely correlated if morphologically linked However the strongly correlated

changes noted by Givnish et al (2005) are ecologically linked but are presumably

morphologicallydevelopmentally independent When there seem to be characters

evolving more or less together tests can be carried out to see if the changes are

concentrated on certain branches of the tree (eg Maddison 1990 Sanderson 1991

Maddison amp Maddison 2000) Such tests have rarely been carried out at higher levels in

angiosperms

One of the most striking examples of a tendency is the distribution of N-fixation

restricted as it is to a monophyletic group of four clades although it has arisen seven (or

perhaps several more) times independently within the N-fixing clade and several

members of two quite different kinds of bacteria are involved (eg Clawson et al 2004

Elliott et al 2007 Sprent amp James 2007) Other examples of work that bears on the issue

of tendencies include the findings that flowers of polysymmetrical Arabidopsis have

genes like TCP1 that are expressed asymmetrically during early development and TCP1

is a probable orthologue of the well-known CYC gene of Antirrhinum that is involved in

the development of monosymmetric slowers there (Cubas et al 2001 Costa et al 2005

etc) Parallelism might build on this underlying morphologically cryptic monosymmetry

even if details of the genetic mechanisms causing the monosymmetry evident in

particular groups may be different (Reeves amp Olmstead 2003 Cubas 2004 see above)

Similarly there have been several recent suggestions that the capability to synthesise a

particular metabolite may be switched off but not lost and so can sometimes be

reacquired (eg Wink amp Witte 1983 Wink 2003 Liscombe et al 2005 Larsson 2007

Waterman 2007) Hence perhaps the rather spotty distribution of many secondary

metabolites like ellagic acid the indole alkaloid camptothecin iridoids etc when

considered in the context of phylogenies As a non-botanical example - but a rather nice

one - Salwini-Plawen and Mayr (1961) suggested some time ago that there has been

considerable parallelism (40-65 or more independent origins) in the evolution of eyes in

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

29

metazoans However Pax 6 seems to be a master control gene that is involved in eye

formation perhaps throughout all bilateralians (eg Gehring amp Ikeo 1999 Erwin amp

Davidson 2002) which changes the nature of the problem Note however that in

general our invocation of evolutionary tendencies to explain the patterns of

distributions of characters is really an admission that we do not understand those

patterns

As we find out more about variation we will probably find fewer and fewer features

constant throughout a group Most unqualified statements of presence and absence

should properly be qualified as usually present or usually absent if one is thinking of

the characterisations on the order pages as encompassing the total variation within a

clade Thus Pistia alone among monocots as so far known has sieve tube plastids with

starch grains not protein crystals However this does not affect the fact that sieve tube

plastids with cuneate protein crystals and lacking starch grains are an apomorphy for

monocots Furthermore fossils in a number of cases suggest character combinations

unknown in extant taxa as may be seen in the discussions of Fagaceae Platanaceae

Iteaceae Calycanthaceae etc Confusing the issue there may be questions as to where

exactly on the tree a particular fossil is to be placed (eg see Nymphaeaceae

Calycanthaceae Archaefructus etc)

To summarize given our current understandings of both phylogenies and

characters evolution of some characters in which we are interested seems very labile

(see eg D Soltis et al 2005b Endress and Matthews 2006a Stevens 2006b Ekman et

al 2008 for a good recent study) and I have been cautious when talking about character

evolution Much effort must continue to be spent in summarizing characters of well-

established clades at all levels providing features by which they may be recognized and

signaling synapomorphies Remember that (1) the basic morphological anatomical and

chemical knowledge of many critical taxa is woefully incomplete (2) different

assumptions about character evolution may greatly affect the position of

synapomorphies on trees (3) in many cases relationships within and between many

groups are too uncertain at present to worry very much about synapomorphies and

(4) we must be clear about what we do and do not not know As mentioned above I

have indicated in a number of places where there are particularly important

uncertainties as to where characters should be placed on the tree Nevertheless it is a

relatively easy matter to update notes such as these and it can be a simple matter to

incorporate new data on characters that have never before been considered in the

context of a tree Much basic - and unfortunately perhaps unfashionable - work must be

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

30

carried to clarify the distribution of morphological anatomical and chemical characters

for examples of what can to be done see the work of P K Endress and collaborators

(floral morphology and development) and S R Jensen and collaborators (iridoids) But

acquiring information about nodal anatomy cell and tissue distribution nodal anatomy

and the like is not difficult at all and should be generally encouraged

But all this is merely a necesary prelude to the understanding of evolution For this

one needs to know a lot more including dating the phylogeny understanding the

function(s) of characters factoring in relevant aspects of the palaeoenvironment in

which an apomorphy first appeared etc

SUMMARY OF THE SYSTEM

Below is a formalised summary of the relationships within orders of the families of seed

plants There are a few families that are not recognised even as options in APG II and

vice versa as well as a few extra orders The families to be recognised in the most

recent edition of Mabberleys The Plant Book (Mabberley 2008) are also largely

consistent with those below But all differences are trivial and will - I hope - eventually

disappear

Square brackets - [] - enclose clades the plus sign - + - designates sister taxa a

comma - - denotes part of a polytomy and quotation marks - - denotes a

paraphyletic group

SEED PLANTS

GYMNOSPERMS

Cycadales

Cycadaceae + Zamiaceae

Ginkgoales

Ginkgoaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

31

Pinales

Pinaceae [[Araucariaceae [Podocarpaceae + Phyllocladaceae]] [Sciadopityaceae

[Taxaceae + Cupressaceae]]]

Gnetales

Ephedraceae [Gnetaceae + Welwitschiaceae]

ANGIOSPERMSFLOWERING PLANTS

Amborellales

Amborellaceae

Nymphaeales

Hydatellaceae [Cabombaceae + Nymphaeaceae]

Austrobaileyales

Austrobaileyaceae [Schisandraceae + Trimeniaceae]

Chloranthales

Chloranthaceae

MAGNOLIIDS

Magnoliales

Myristicaceae [Magnoliaceae [[Himantandraceae + Degeneriaceae] [Eupomatiaceae +

Annonaceae]]]

Laurales

Calycanthaceae [[Siparunaceae [Gomortegaceae + Atherospermataceae]] [Monimiaceae

[Hernandiaceae + Lauraceae]]]

Canellales

Canellaceae + Winteraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

32

Piperales

[Hydnoraceae + Aristolochiaceae] [Piperaceae + Saururaceae]

MONOCOTS

Acorales

Acoraceae

Alismatales

Araceae [Tofieldiaceae [[Alismataceae [Hydrocharitaceae + Butomaceae]]

[Scheuchzeriaceae [Aponogetonaceae [Juncaginaceae [Maundiaceae [[Posidoniaceae

[Ruppiaceae + Cymodoceaceae]] [Zosteraceae + Potamogetonaceae]]]]]]]

Petrosaviales

Petrosaviaceae

Dioscoreales

Nartheciaceae [[Taccaceae + Thismiaceae] [Burmanniaceae + Dioscoreaceae]]

Pandanales

Velloziaceae Triuridaceae Stemonaceae [Pandanaceae + Cyclanthaceae]

Liliales

Corsiaceae [Campynemataceae [Petermanniaceae [Colchicaceae + Alstroemeriaceae]]

Melanthiaceae [[Philesiaceae + Rhipogonaceae] [Smilacaceae + Liliaceae]]]

Asparagales

Orchidaceae [[Boryaceae [Blandfordiaceae [Lanariaceae [Asteliaceae + Hypoxidaceae]]]]

[[Ixioliriaceae + Tecophilaeaceae] [Doryanthaceae [Iridaceae [Xeronemataceae

[Xanthorrhoeaceae [Alliaceae + Asparagaceae]]]]]]]

COMMELINIDS

Unplaced

Dasypogonaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

33

Arecales

Arecaceae

Poales

[Typhaceae + Bromeliaceae] [Rapateaceae [[Thurniaceae [Juncaceae + Cyperaceae]]

[[Mayacaceae [Eriocaulaceae + Xyridaceae]] [Flagellariaceae [Anarthriaceae

[Centrolepidaceae + Restionaceae]]] [Joinvilleaceae [Ecdeiocoleaceae + Poaceae]]]]]]]

Commelinales

[Commelinaceae + Hanguanaceae] [Philydraceae [Haemodoraceae + Pontederiaceae]]

Zingiberales

Musaceae [Strelitziaceae + Lowiaceae] Heliconiaceae [[Cannaceae + Marantaceae]

[Costaceae + Zingiberaceae]]

Ceratophyllales

Ceratophyllaceae

EUDICOTS

Ranunculales

Eupteleaceae [Papaveraceae [[[Lardizabalaceae + Circaeasteraceae] [Menispermaceae

[Berberidaceae + Ranunculaceae]]]]

Sabiales

Sabiaceae

Proteales

Nelumbonaceae [Platanaceae + Proteaceae]

Trochodendrales

Trochodendraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

34

Buxales

Haptanthaceae [Buxaceae + Didymelaceae]

CORE EUDICOTS

Gunnerales

Gunneraceae + Myrothamnaceae

Dilleniales

Dilleniaceae

Saxifragales

Peridiscaceae [[Paeoniaceae [Altingiaceae [Hamamelidaceae [Cercidiphyllaceae +

Daphniphyllaceae]]]] [[Crassulaceae [Aphanopetalaceae [Tetracarpaeaceae

[Penthoraceae + Haloragaceae]]]] [Iteaceae [Grossulariaceae + Saxifragaceae]]]]

Cynomoriaceae unplaced

Vitales

Vitaceae

ROSIDS

FABIDROSID I

Zygophyllales

Krameriaceae + Zygophyllaceae

Celastrales

Lepidobotryaceae + Celastraceae

Oxalidales

Huaceae [[Connaraceae + Oxalidaceae] [Cunoniaceae [Elaeocarpaceae [Brunelliaceae +

Cephalotaceae]]]]

Malpighiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

35

[Achariaceae [Goupiaceae [Violaceae + Passifloraceae] [Lacistemataceae Salicaceae]]

[Lophopyxidaceae Putranjivaceae] [Ctenolophonaceae [Erythroxylaceae +

Rhizophoraceae]] Linaceae Ixonanthaceae Humiriaceae Irvingiaceae

Centroplacaceae Pandaceae [[Bonnetiaceae + Clusiaceae] [Calophyllaceae

[Hypericaceae + Podostemaceae]]] [Malpighiaceae + Elatinaceae] Ochnaceae

[Peraceae [Rafflesiaceae + Euphorbiaceae]] [Phyllanthaceae + Picrodendraceae]

[Balanopaceae [[Trigoniaceae + Dichapetalaceae] [Chrysobalanaceae +

Euphroniaceae]]] Caryocaraceae

N-FIXING CLADE

Fabales

Quillajaceae [Fabaceae [Polygalaceae + Surianaceae]]

Rosales

Rosaceae [[Barbeyaceae [Elaeagnaceae Dirachmaceae Rhamnaceae]] [Ulmaceae

[Cannabaceae [Moraceae + Urticaceae]]]]

Cucurbitales

Anisophylleaceae [[Corynocarpaceae + Coriariaceae] [Cucurbitaceae [Tetramelaceae

[Datiscaceae + Begoniaceae]]]] Apodanthaceae

Fagales

Nothofagaceae [Fagaceae [[Myricaceae + Juglandaceae] [Casuarinaceae

[Ticodendraceae + Betulaceae]]]]

MALVIDROSID II

pgtGeraniales

Geraniaceae [[Melianthaceae + Francoaceae] [Vivianaceae + Ledocarpaceae]]

Myrtales

Combretaceae [[Onagraceae + Lythraceae] [[Vochysiaceae + Myrtaceae]

[Melastomataceae [Crypteroniaceae [Alzateaceae + Penaeaceae]]]]]

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

36

Crossosomatales

[Staphyleaceae [Guamatelaceae [Crossosomataceae + Stachyuraceae]]] [Aphloiaceae

[Geissolomataceae + Strasburgeriaceae]]

Picramniales

Picramniaceae

Sapindales

Biebersteiniaceae Nitrariaceae [Kirkiaceae [Anacardiaceae + Burseraceae]]

[Sapindaceae [Simaroubaceae Rutaceae Meliaceae]]

Huerteales

Gerradinaceae [Dipentodontaceae + Tapisciaceae]

Malvales

Neuradaceae [Thymelaeaceae [Sphaerosepalaceae Bixaceae [Cistaceae

[Sarcolaenaceae + Dipterocarpaceae]] [Cytinaceae + Muntingiaceae] Malvaceae]]

Brassicales

[Akaniaceae + Tropaeolaceae] [[Moringaceae + Caricaceae] [Setchellanthaceae

[Limnanthaceae [[Koeberliniaceae [Bataceae + Salvadoraceae]] [Emblingiaceae

[Pentadiplandraceae [Gyrostemonaceae Resedaceae Stixis and relatives] Tovariaceae

[Capparaceae [Cleomaceae + Brassicaceae]]]]]]]]

Berberidopsidales

Aextoxicaceae + Berberidopsidaceae

Santalales

Erythropalaceae Olacaceae [[Loranthaceae [Misodendraceae + Schoepfiaceae]]

[Opiliaceae + Santalaceae]] Balanophoraceae unplaced

Caryophyllales

[[Droseraceae [Nepenthaceae [Drosophyllaceae [Ancistrocladaceae +

Dioncophylleaceae]]]]] [[Frankeniaceae + Tamaricaceae] [Polygonaceae +

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

37

Plumbaginaceae]]] [Rhabdodendraceae [Simmondsiaceae [[Asteropeiaceae +

Physenaceae] [[Caryophyllaceae [Achatocarpaceae + Amaranthaceae]]

[Stegnospermataceae [Limeaceae [[Lophocarpaceae [Barbeuiaceae [Aizoaceae

[Sarcobataceae Phytolaccaceae Nyctaginaceae]]]] [Molluginaceae [Halophytaceae

Montiaceae Basellaceae Didiereaceae [Talinaceae [Portulacaceae [Anacampseros etc +

Cactaceae]]]]]]]]]]]]]

ASTERIDS

Cornales

[Cornaceae + Nyssaceae] [Hydrangeaceae + Loasaceae] Hydrostachyaceae [Curtisiaceae

+ Grubbiaceae]

Ericales

[Balsaminaceae [Marcgraviaceae + Tetrameristaceae]] [[Polemoniaceae +

Fouquieraceae] Lecythidaceae [[Sladeniaceae + Pentaphylacaceae] [Sapotaceae

[Ebenaceae [Maesaceae [Theophrastaceae [Myrsinaceae + Primulaceae]]]]]

[Mitrastemonaceae Theaceae [Symplocaceae [Styracaceae + Diapensiaceae]]

[[Sarraceniaceae [Roridulaceae + Actinidiaceae] [Clethraceae [Cyrillaceae +

Ericaceae]]]]]]

LAMIIDASTERID I

Unplaced

Metteniusaceae Oncothecaceae Icacinaceae and some unplaced ex-Icacinaceae

Garryales

Garryaceae + Eucommiaceae

Unplaced

Boraginaceae Vahliaceae

Gentianales

Rubiaceae [Gentianaceae [Loganiaceae [Gelsemiaceae + Apocynaceae]]]

Lamiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

38

Plocospermataceae [Oleaceae [Tetrachondraceae [Calceolariaceae

[Gesneriaceae Peltanthera [Pedaliaceae Carlemanniaceae Martyniaceae Stilbaceae

Plantaginaceae Linderniaceae Bignoniaceae Verbenaceae Lamiaceae Paulowniaceae

Schlegeliaceae Thomandersiaceae Phrymaceae Rehmannia Acanthaceae

Scrophulariaceae Orobanchaceae Byblidaceae Lentibulariaceae]]]]]

Solanales

[Montiniaceae [Sphenocleaceae + Hydroleaceae]] [Convolvulaceae + Solanaceae]

CAMPANULIDASTERID II

Aquifoliales

[[Cardiopteridaceae + Stemonuraceae] [Aquifoliaceae [Helwingiaceae +

Phyllonomaceae]]

Asterales

[Rousseaceae + Campanulaceae] Pentaphragmataceae [[Alseuosmiaceae [Phellinaceae

+ Argophyllaceae]] Stylidiaceae [Menyanthaceae [Goodeniaceae [Calyceraceae +

Asteraceae]]]]

Escalloniales

Escalloniaceae

Bruniales

[Bruniaceae + Columelliaceae]

Apiales

Pennantiaceae [Torricelliaceae [Griseliniaceae [Pittosporaceae [Araliaceae

[Myodocarpaceae + Apiaceae]]]]]

Paracryphiales

Paracryphiaceae

Dipsacales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

39

Adoxaceae [Diervillaceae [Caprifoliaceae [Linnaeaceae [Morinaceae [Dipsacaceae +

Valerianaceae]]]]]

LINKS TO ORDERS AND FAMILIES

[Back to Top]

MAIN GROUPINGS

Asterids commelinids core eudicots asterid 1 asterid

2 eudicots gymnosperms Magnoliophyta monocots N-fixing claderosids seed plants

ALPHABETICAL LISTING OF ALL ORDINAL NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | L | M | N | O | P | Q | R | S | T | U | V | W | X | Z

Abietales Acanthales Acerales Acorales Actinidiales Actinostrobales Adoxales Aescul

ales Agavales Aizoales AkanialesAlismatales Alliales Alseuosmiales Alstroemeriales

Altingiales Amaranthales Amaryllidales Amborellales AmbrosialesAmmiales Amomal

es Ancistrocladales Anisophylleales Annonales Anthobolales Apiales Apocynales Apo

nogetonalesAquifoliales Arales Araliales Aralidiales Araucariales Arecales Aristolochi

ales Athrotaxidales Asarales AsclepiadalesAsparagales Asphodelales Asparagales Ast

eliales Atriplicales Aucubales Austrobaileyales Avenales

Balanitales Balanopales Balanophorales Balsaminales Barbeyales Barclayales Batales

Begoniales BerberidalesBerberidopsidales Betulales Biebersteiniales Bignoniales Bix

ales Boraginales Brassicales Brexiales Bromeliales BrunialesBrunoniales Burmannial

es Burserales Butomales Buxales Byblidales

Cactales Callitrichales Calycanthales Calycerales Campanulales Campynematales Can

ellales Cannales CapparalesCaprifoliales Cardiopteridales Carduales Caricales Carle

manniales Caryophyllales Cassiales Casuarinales CelastralesCentrolepidales Cephalot

ales Cephalotaxales Ceratophyllales Cercidiphyllales Chenopodiales Chironiales Chlor

anthalesChrysobalanales Cinchonales Circaeasterales Cistales Citrales Cocosales Col

chicales Columelliales CombretalesCommelinales Connarales Convolvulales Coriarial

es Cornales Corylales Corynocarpales Crassulales CrossosomatalesCucurbitales Cunn

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

40

inghamiales Cunoniales Cupressales Cycadales Cyclanthales Cymodoceales Cynarales

CynomorialesCyperales Cyrillales Cytinales

Daphnales Daphniphyllales Dasypogonales Datiscales Degeneriales Desfontainiales D

iapensiales Didymelales DillenialesDioales Dioncophyllales Dioscoreales Diospyrales

Dipentodontales Dipsacales Droserales

Ebenales Echiales Elaeagnales Elaeocarpales Elatinales Elodeales Emmotales Empetr

ales Ephedrales Ericales EriocaulalesErythropalaless Escalloniales Eucommiales Euph

orbiales Eupomatiales Eupteleales Euryalaless

Fabales Fagales Falcatifoliales Ficales Flacourtiales Flagellariales Fouquieriales Franc

oales Frangulales

Galiales Garryales Geissolomatales Gentianales Geraniales Gesneriales Ginkgoales G

laucidiales Globulariales GnetalesGoodeniales Greyiales Griseliniales Grossulariales

Grubbiales Gunnerales Gyrocarpales Gyrostemonales

Haemodorales Haloragales Hamamelidales Hanguanales Heisteriales Helleborales He

lwingiales Himantandrales HippuridalesHomaliales Hortensiales Huales Huerteales

Hydatellales Hydnorales Hydrangeales Hydrastidales HydrocharitalesHydropeltidales

Hydrostachyales Hypericales Hypoxidales

Icacinales Illiciales Iridales Irvingales Iteales Ixiales

Jasminales Juglandales Julianiales Juncaginales Juncales

Lacistematales Lactoridales Lamiales Lardizabalales Laurales Lecythidales Ledocarpal

es Leitneriales LentibularialesLigustrales Liliales Limnanthales Linales Liriales Loasal

es Lobeliales Loganiales Lonicerales Loranthales LowialesLythrales

Magnoliales Malpighiales Malvales Marathrales Marcgraviales Mayacales Medusagy

nales Medusandrales MelanthialesMelastomatales Meliales Melianthales Meliosmal

es Menispermales Menyanthales Metteniusales MitrastemonalesMiyoshiales Moni

miales Moringales Musales Myricales Myristicales Myrothamnales Myrsinales Myrta

les

Najadales Nandinales Narcissales Nartheciales Nelumbonales Nepenthales Neuradal

es Nitrariales Nolanales NothofagalesNyctaginales Nymphaeales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

41

Ochnales Oenotherales Olacales Oleales Onagrales Oncothecales Opuntiales Orchid

ales Oxalidales

Saxifragales Pandales Pandanales Papaverales Paracryphiales Parasitaxales Paridales

Parnassiales PassifloralesPenaeales Pennantiales Pentaphragmatales Peridiscales P

etiveriales Petrosaviales Phellinales Philydrales PhyllanthalesPhysenales Phytolaccal

es Picramniales Pinales Pinguiculales Piperales Pittosporales Plantaginales Platanale

sPlumbaginales Poales Podocarpales Podophyllales Podostemales Polemoniales Pol

ygalales Polygonales PontederialesPortulacales Posidoniales Potamogetonales Primu

lales Proteales

Quercales Quillajales Quintiniales

Rafflesiales Ranunculales Rapateales Resedales Restionales Rhabdodendrales Rhamn

ales Rhinanthales RhizophoralesRhodorales Rhoipteleales Roridulales Rosales Rouss

eales Rubiales Ruppiales Rutales

Sabiales Salicales Salvadorales Samolales Samydales Sanguisorbales Santalales Sapin

dales Sapotales SarracenialesSaxegotheales Saxifragales Scheuchzeriales Sciadopitya

les Scleranthales Scrophulariales Scyphostegiales SedalesSilenales Simmondsiales S

milacales Solanales Sphenocleales Sphenostemonales Stangeriales Stellariales Stemo

nalesStilbales Stylidiales Styracales Surianales

Taccales Tamales Tamaricales Taxales Taxodiales Tecophilaeales Terebinthales Tern

stroemiales Theales TheligonalesThymelaeales Tiliales Tofieldiales Torricelliales Tov

ariales Tribelales Trilliales Trimeniales Triuridales TrochodendralesTropaeolales Turn

erales Typhales

Ulmales Urticales

Vacciniales Vahliales Vallisneriales Velloziales Veratrales Verbenales Viburnales Vinc

ales Violales Viscales VitalesVochysiales

Welwitschiales Winterales

Xanthorrhoeales Ximeniales Xyridales

Zamiales Zingiberales Zosterales Zygophyllales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

42

ALPHABETICAL LISTING OF ALL FAMILY NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | K | L | M | N | O | P | Q | R | S | T | U | V | W | X | Y

|Z

Back to Home

In some cases clicking on a family synonym will send you directly to the subfamily in

which the genus of the basionym is to be found

Abietaceae Abolbodaceae Abrophyllaceae Acalyphaceae Acanthaceae [Acanthochla

mydaceae] Acarnaceaee AceraceaeAchariaceae Achatocarpaceae Achradaceae Achy

ranthaceae Acmopylaceae Acoraceae Acristaceae ActaeaceaeActinidiaceae Actinota

ceae Amentotaxaceae Adenogrammaceae Adoxaceae Aegialitidaceae Aegicerataceae

Aegilopaceae Aeginetiaceaee Aegiphilaceae Aesculaceae Aextoxicaceae Agapanthac

eae Agavaceae AgdestidaceaeAgialidaceae Agrimoniaceae Agrostidaceae Ailanthace

ae Aitoniaceae Aizoaceae Akaniaceae Alangiaceae AlchemillaceaeAldrovandaceae A

lismataceae Alliaceae Allioniaceae Allophylaceae Aloaceae Alopecuraceae Alpiniacea

e AlseuosmiaceaeAlsinaceae Alsodeiaceae Alstroemeriaceae Altingiaceae Alzateacea

e Amaranthaceae Amaryllidaceae Amborellaceae Ambrosiaceae Amentotaxaceae A

mmanniaceae Ammiaceae Amomaceae Ampelidopsaceae Amygdalaceae Amyridacea

eAnacardiaceae Anagallidaceae Anarthriaceae Anchusaceae Ancistrocladaceae Andr

omedaceae AndropogonaceaeAndrostachyaceae Androsynaceae Anemarrhenaceae

Anemonaceae Angelicaceae Anisophylleaceae AnnonaceaeAnomochloaceae Anopter

aceae Anrederaceae Anthemidaceae Anthericaceae Anthobolaceae Antidesmataceae

Antirrhinaceae Antoniaceae Aparinaceae Aphanopetalaceae Aphloiaceae Aphyllanth

aceae Apiaceae ApocynaceaeApodanthaceae Aponogetonaceae Aporusaceae Apose

ridaceae Apostasiaceae Aptandraceae AquifoliaceaeAquilariaceae Aquilegiaceae A

raceae Aragoaceae Araliaceae Aralidiaceaee Araucariaceae Arbutaceae Arceuthidace

aeArceuthobiaceae Arctostaphylaceae Arctotidaceae Ardisiaceae Arecaceae Argoph

yllaceae Arisaraceae AristolochiaceaeAristoteliaceae Arjonaceae Armeriaceae Artem

isiaceae Arthrotaxidaceae Artocarpaceae Arundinaceae ArundinellaceaeAsaraceae A

sclepiadaceae Ascyraceaee Asiraceae Asparagaceae Asperulaceae Asphodelaceae As

pidistraceaeeAsteliaceae Asteraceae Asteranthaceae Asteropeiaceae Astragalaceae

Astrocarpaceae AthanasiaceaeAtherospermataceae Atriplicaceae Atropaceae Aucuba

ceae Aurantiaceae Austrobaileyaceae Austrotaxaceae AvenaceaeAverrhoaceae Avet

raceae Avicenniaceae Azaleaceae Azimaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

43

Balanitaceae Balanopaceae Balanophoraceae Balsameaceae Balsaminaceae Bambusa

ceae Barbaceniaceae BarbeuiaceaeBarbeyaceae Barclayaceae Barringtoniaceae Base

llaceae Bataceae Baueraceae Bauhiniaceae Baxteriaceae BegoniaceaeBehniaceae B

elangeraceae Belloniaceae Bembiciaceae Berberidaceae Berberidopsidaceae Berryac

eaee BersamaceaeBertyaceae Berzeliaceae Besleriaceae Betaceae Betulaceae Biebe

rsteiniaceae Bifariaceae Bignoniaceae BischofiaceaeBixaceae Blakeaceae Blandfordia

ceae Blattiaceae Blepharocaryaceae Blitaceae Blyxaceae Boerlagellaceae Bolivaracea

eBombacaceae Bonnetiaceae Bontiaceae Boopidaceae Boraginaceae Borassaceae B

oroniaceae BoryaceaeBotryodendraceae Bougainvilleaceae Boweniaceae Brassicacea

e Brachycaulaceae Bretschneideraceae BrexiaceaeBromeliaceae Brownlowiaceae Br

unelliaceae Bruniaceae Brunoniaceae Brunsvigiaceae Bryoniaceae BuchneraceaeBuci

daceae Buddlejaceae Buglossaceae Bulbocodiaceae Bumeliaceae Bupleuraceae Burc

hardiaceae BurmanniaceaeBurseraceae Butneriaceae Butomaceae Buxaceae Byblida

ceae Byttneriaceae

Cabombaceae Cacaoaceae Cactaceae Caesalpiniaceae Caladiaceae Calamaceae Calce

olariaceae CalectasiaceaeCalendulaceae Callaceae Callicomaceae Calligonaceae Calli

traceae Callitrichaceae Calochortaceae CalophyllaceaeCalthaceae Calycanthaceae Ca

lyceraceae Cambogiaceae Camelliaceae Campanulaceae CampynemataceaeCanacom

yricaceae Canellaceae Cannabaceae Cannaceae Canopodaceae Canotiaceae Cansjera

ceae CapparaceaeCaprariaceae Caprifoliaceae Cardiopteridaceae Carduaceae Carica

ceae Carissaceae Carlemanniaceae CarpinaceaeCarpodetaceae Cartonemataceae Ca

ryocaraceae Caryophyllaceae Casearieaceae Cassiaceae CassipoureaceaeCassythacea

e Castaneaceae Castelaceae Casuarinaceae Catesbaeaceae Cathedraceae Cecropiace

ae CedraceaeCedrelaceae Celastraceae Celosiaceae Celtidaceae Centaureaceae Cen

trolepidaceae Centroplacaceae CepaceaeCephalanthaceae Cephalotaceae Cephalota

xaceae Cerastiaceae Ceratoniaceae Ceratophyllaceae CerberaceaeCercidiphyllaceae

Cercocarpaceae Cercodiaceae Cereaceae Cerinthaceae Ceroxylaceae Cestraceae Cev

alliaceaeChailletiaceae Chamaedoraceae Chamaemoraceae Chamelauciaceae Chaun

ochitonaceae Chelidoniaceae ChelonaceaeChenopodiaceae Chimonanthaceae Chingi

thamnaceae Chionographidaceae Chiranthodendraceae ChironiaceaeChlaenaceae Ch

loanthaceae Chloranthaceae Chloridaceae Chlorogalaceae Chrysobalanaceae Cicerac

eae CichoriaceaeCimicifugaceae Cinchoniaceae Circaeaceae Circaeasteraceae Cissac

eae Cistaceae Citraceae Clematidaceae CleomaceaeClethraceae Clusiaceae Cneorac

eae Cnestidaceae Cnicaceae Cobaeaceae Cochlospermaceae Cocosaceae Coffeaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Colchicaceae Coleogynaceae Columelliaceae Combretaceae Commelinaceae Comocl

adiaceae Compositae CompsoaceaeConantheraceae Connaraceae Conostylidaceae C

onvallariaceae Convolvulaceae Cordiaceae CoreopsidaceaeCoriandraceae Coriariacea

e Coridaceae Corispermaceae Cornaceae Corokiaceae Coronillaceae Correaceae Cor

rigiolaceaeCorsiaceae Corydalaceae Corylaceae Corynocarpaceae Coryphaceae Cost

aceae Cotyledonaceae CoulaceaeCoutariaceae Coutoubiaceae Crassulaceae Crescen

tiaceae Cressaceae Crinaceae Crocaceae CroomiaceaeCrossosomataceae Crotonacea

e Cruciferae Cryptaceae Crypteroniaceae Cryptocorynaceae CryptomeriaceaeCtenol

ophonaceae Cucurbitaceae Cunninghamiaceae Cunoniaceae Cupressaceae Curcumac

eae Curtisiaceae CuscutaceaeCyananthaceae Cyanastraceae Cynanchaceae Cyanella

ceae Cycadaceae Cyclanthaceae CyclantheraceaeCyclocheilaceae Cydoniaceae Cymo

doceaceae Cynaraceae Cynocrambaceae Cynomoriaceae Cyperaceae CyphiaceaeCyp

hocarpaceae Cypripediaceae Cyrillaceae Cyrtandraceae Cyrtanthaceae Cytinaceae

Dacrycarpaceae Dactylanthaceae Damasoniaceae Daphnaceae Daphniphyllaceae Das

ypogonaceae DatiscaceaeDaturaceae Daucaceae Davidiaceae Davidsoniaceae Decais

neaceae Deeringiaceae Degeneriaceae DelphiniaceaeDendrophthoaceae Columelliac

eae Detariaceae Dialypetalanthaceae Dianellaceae Dianthaceae DiapensiaceaeDicha

petalaceae Dichondraceae Diclidantheraceae Dicrastylidiaceae Dictamnaceae Didiere

aceae DidymelaceaeDidymocarpaceae Diegodendraceae Diervillaceae Digitalidaceae

e Dilatridaceae Dilleniaceae Dioaceae DionaeaceaeDioncophyllaceae Dioscoreaceae

Diosmaceae Diospyraceae Dipentodontaceae Diphylleiaceae Diplolaenaceae Dipsaca

ceaeDipterocarpaceae Dirachmaceae Disanthaceae Diselmaceae Dodonaeaceae Do

mbeyaceae Donatiaceae DorsteniaceaeDortmannaceae Doryanthaceae Drabaceae D

racaenaceae Dracontiaceae Drimyidaceae Droseraceae DrosophyllaceaeDryadaceae

Duabangaceae Duckeodendraceae Dulongiaceae Durantaceae Durionaceae Dysphani

aceae

Ebenaceae Eccremidaceae Ecdeiocoleaceae Echiaceae Echinopaceae Ehretiaceae Ela

eagnaceae ElaeocarpaceaeElegiaceae Elatinaceae Ellisiaceae Ellisiophyllaceae Elodea

ceae Elytranthaceae Embeliaceae Emblingiaceae EmmotaceaeEmpetraceae Encepha

lartaceae Engelhardtiaceae Enhalaceae Epacridaceae Ephedraceae Ephemeranthacea

eEpilobiaceae Epimediaceae Eragrostidaceae Eremolepidaceae Eremosynaceae Erica

ceae Erinaceae EriocaulaceaeEriogonaceae Eriospermaceae Erodiaceae Erycibaceae

Eryngiaceae Erysimaceae Erythroniaceae ErythropalaceaeErythrospermaceae Erythro

xylaceae Escalloniaceae Eschscholziaceae Eucomidaceae Eucommiaceae Eucryphiace

aeEuonymaceae Eupatoriaceae Euphorbiaceae Euphrasiaceae Euphroniaceae Eupo

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

45

matiaceae Eupteleaceae EuryalaceaeEustrephaceae Euthemidaceae Eutocaceae Exb

ucklandiaceae Exocarpaceae

Fabaceae Fagaceae Falcatifoliaceae Ferulaceae Festucaceae Ficaceae Ficoidaceae Fit

zroyaceae FlacourtiaceaeFlagellariaceae Flindersiaceae Foetidiaceae Forestieraceae

Fothergillaceae Fouquieriaceae Fragariaceae FrancoaceaeFrangulaceae Frankeniacea

e Fraxinaceae Fraxinellaceae Freycinetiaceae Fritillariaceae Fuchsiaceae Fumariacea

eFunkiaceae

Gaiadendraceae Galacaceae Galanthaceae Galaxiaceae Galedupaceae Galeniaceae G

aliaceae Garciniaceae GardeniaceaeGardneriaceae Garryaceae Geissolomataceae Ge

itonoplesiaceae Gelsemiaceae Geniostomaceae GentianaceaeGeonomataceae Geosir

idaceae Geraniaceae Gerrardinaceae Gesneriaceae Gethyllidaceae Gilliesiaceae Gina

lloaceaeGinkgoaceae Gisekiaceae Gladiolaceae Glaucidiaceae Glechomaceae Glinace

ae Globulariaceae Gnaphaliaceae GnetaceaeGoetzeaceae Gomortegaceae Gomphiac

eae Gomphrenaceae Gonystylaceae Goodeniaceae Gordoniaceae GouaniaceaeGoup

iaceae Gramineae Gratiolaceae Grewiaceae Greyiaceae Grielaceae Griseliniaceae Gr

onoviaceae GrossulariaceaeGrubbiaceae Guaiacanaceae Guamatelaceae Guettardac

eae Gunneraceae Gustaviaceae Guttiferae GyrocarpaceaeGyrostemonaceae

Hachetteaceae Haemanthaceae Haemodoraceae Halesiaceae Hallieraceae Halocarpa

ceae Halophilaceae HalophytaceaeHaloragaceae Hamamelidaceae Hanguanaceae H

aptanthaceae Harmandaceae Hebenstretiaceae HectorellaceaeHederaceae Hedyosm

aceae Hedyotidaceae Hedysaraceae Heisteriaceae Heleniaceae Heliamphoraceae H

elianthaceaeHelianthemaceae Helichrysaceae Heliconiaceae Helicteraceae Heliotropi

aceae Helleboraceae Heloniadaceae HelosidaceaeHelwingiaceae Hemerocallidaceae

Hemimeridaceae Hemiphylacaceae Henriqueziaceae Henslowiaceae Hermanniaceae

Hernandiaceae Herniariaceae Herreriaceae Hesperocallidaceae Heterantheraceae He

teropyxidaceae HeterostylaceaeHibbertiaceae Hibiscaceae Hilleriaceae Himantandra

ceae Hippocastanaceae Hippocrateaceae HippomanaceaeHippophaeaceae Hippurida

ceae Hirtellaceae Holacanthaceae Homaliaceae Hopkinsiaceae Hoplestigmataceae H

ordeaceaeHornschuchiaceae Hortensiaceae Hortoniaceae Hostaceae Houstoniaceae

Huaceae Huerteaceae HugoniaceaeHumbertiaceae Humiriaceae Hyacinthaceae Hyd

atellaceae Hydnoraceae Hydrangeaceae Hydrastidaceae HydrillaceaeHydrocharitacea

e Hydrocotylaceae Hydrogetonaceae Hydroleaceae Hydropeltidaceae Hydrophylacac

eae HydrophyllaceaeHydrostachyaceae Hymenocardiaceae Hypecoaceae Hyperanth

eraceae Hypericaceae Hypopityaceae HypoxidaceaeHypseocharitaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

46

Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta

ceae ImperatoriaceaeInocarpaceae Inulaceae Ionidiaceae Iriarteaceae Iridaceae Irvi

ngiaceae Isnardiaceae Isophysidaceae IteaceaeStrasburgeriaceae Ixiaceae Ixioliriace

ae Ixonanthaceae

Jalapaceae Jamboliferaceae Japonoliriaceae Jasionaceae Jasminiaceae Johnsoniaceae

Joinvilleaceae JuglandaceaeJulianiaceae Juncaceae Juncaginaceae Juniperaceae Jus

siaeaceae Justiciaceae

Kadsuraceae Kaliphoraceae Kalmiaceae Kaniaceae Kiggelariaceae Kingdoniaceae Kin

giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae

Krameriaceae

Labiatae Lacandoniaceae Lachenaliaceae Lacistemataceae Lactoridaceae Lactucaceae

Lagerstroemiaceaee LamiaceaeLampsanaceae Lanariaceae Langsdorffiaceae Lantan

aceae Lapageriaceae Lardizabalaceae Lasiopetalaceae LauraceaeLawsoniaceae Laxm

anniaceae Lecythidaceae Ledaceae Ledocarpaceae Leeaceae Leguminosae Leitneriac

eae LemnaceaeLennoaceae Lentibulariaceae Lentiscaceae Leoniaceae Leonticaceae

Lepidobotryaceae LepidocarpaceaeLepidocaryaceae Lepidocerataceae Leptaulaceae

Leptospermaceae Lepturaceae Lepuropetalaceae LeuctenbergiaceaeLibocedraceae

Licaniaceae Ligustraceae Lilaeaceae Liliaceae Limeaceae Limnanthaceae Limnocharit

aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace

ae Linderniaceae Lindleyaceae Linnaeaceae LippayaceaeLiriaceae Liriodendraceae Li

ssocarpaceae Littorellaceae Loasaceae Lobeliaceaee Loganiaceae LomandraceaeLoni

ceraceae Lopeziaceae Lophiocarpaceae Lophiolaceae Lophiraceae Lophophytaceae L

ophopyxidaceae LoranthaceaeLotaceae Lowiaceae Lupulaceaee Luxemburgiaceae Lu

zuriagaceae Lychnidaceae Lyciaceae LyginiaceaeLygodisodeaceae Lysimachiaceae Ly

thraceae

Macarisiaceae Mackinlayaceae Madiaceae Maesaceae Magnoliaceae Malaceae Male

sherbiaceae MalortieaceaeMalpighiaceae Malvaceae Mangiaceae Manicariaceae Ma

paniaceae Marantaceae Marathraceae MarcgraviaceaeMartyniaceae Mastixiaceae

Matricariaceae Maundiaceae Mayacaceae Medeolaceae Medusagynaceae Peridiscac

eaeMelaleucaceae Melampyraceae Melanophyllaceae Melanthiaceae Melastomatac

eae Meliaceae Melianthaceae MelicaceaeMeliosmaceae Melittidaceae Melochiacea

e Memecylaceae Mendonciaceae Menispermaceae Menthaceae MenyanthaceaeMe

nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

47

Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

Nypaceae Nyssaceae

Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

ceae Papilionaceae Pappophoraceae PapyraceaeParacryphiaceae Parasitaxaceae Par

ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

48

hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

49

cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

50

ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 10: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

10

etc as is evident in Takhtajans own work - general taxonomic inflation is the result (see

also comparable suggestions in a cladistic context for Brassicales in particular - Ronse de

Craene amp Haston 2006) Such splitting is also questionable when teaching and learning

families since the student needs to understand the system as a whole However for

some genera removed from the families that until now have included them the

phenetic-classificatory-phylogenetic structure in their new home may mandate the

recognition of small families On the other hand Takhtajans suggestion that narrowly

defined families are more useful for phylogenetic studies may be true Indeed I have

more than once regretted prematurely combining groups whether species (in the

context of monographic work) or families (in the course of preparing these notes) but

this is largely a separate issue

Van Steenis (1978) Philipson (1987b) and others have rightly questioned the value

of splitting a group when ideas of the relationships of its constituent members have not

changed - that is very good reasons have to be provided for splitting a family if the

genera within it remain part of the same clade rather than belonging to another clade

Thus APG (2003) broadened the circumscription of Malvaceae because of the

parapolyphyly of some of the families that had historically been associated with it (Judd

amp Manchester 1997 Alverson et al 1999 Bayer et al 1999) These families particularly

Tiliaceae and Sterculiaceae were not at all easy to distinguish their close relationship

(see eg Brown 1814) and overall similarity had long been conceded and to some

workers at least their combination has come as something of a relief Although most of

the larger clades within Malvaceae sl remain difficult to distinguish even with flowers

Cheek (2007) opts for a wholesale and novel dismemberment into ten families

however the very good reasons for doing this are wanting

The same principles are of course applicable when it comes to dividing genera little

other than a headache is gained by splitting genera such

as Drosera and Gnetum (Doweld 2000) as has recently been proposed Thus if an

established genus divides into two (or more) clades this is not a signal for recognising

two groups at the same level - so here it could be argued that the dismemberment

of Pterostylis (Jones amp Clements 2002b) was somewhat unfortunate Along the same

lines if a newly-discovered taxon is sister to an existing named taxon - say a genus - this

does not necessitate the description of a separate genus for the newly described species

(cf Davis 2002) (Of course some [eg Thorne 1976] have suggested that the sizes of

gaps between groups at the same rank should be similar but any principle like this is

inherently flawed since morphological gaps are more unstable than phylogenetic

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

11

relationships and applying it across all flowering plants would both be difficult and cause

substantial changes - see also below) Humphreys and Linder (2009) provide a well-

documented survey of generic concepts in plants which the reader should consult they

note that generic limits (broad versus narrow) have oscillated historically and that

currently larger genera tend to be recognised because studies tend to be on a broader

scale than in the past

Note that invoking similarity or difference - whether qualified (considerable

similarities substantial differences) or not - in a cladistic context as justification for

combining or splitting taxa is not a particularly strong argument (see eg

CardiopteridaceaeStemonuraceae - Karingrehed 2002c) Similarity and difference can

neither be defined precisely since what may seem to be substantial similarities to me

may not to the next person nor are they likely to be stable in the face of our changing

knowledge of morphology and what might be synapomorphies

I might have prefered to merge some families recognised here or split others but

by and large I do not think my own preferences matter very much - and I take the same

position with regards to comparable preferences expressed by others Indeed the

bottom line is that in flagged hierarchies of the kind used here the limits of any

monophyletic unit generally taught and discussed particularly other than species can

be established only by convention and consensus (eg Stevens 2002 2006a Entwisle amp

Weston 2005) This is rather different from the reasonable observation that the

phylogeny itself cannot be achieved by consensus (Thorne 1976) for in most cases there

is a fact of the matter when it comes to relationships Given the increasing support for

the outlines of angiosperm phylogeny a stable consensus classification based on this

phylogeny seems attainable Indeed in addition to providing current ideas of

relationships of seed plants in a synthesised form this site is part of an attempt to build

such a consensus about the circumscription of taxa (see APG 1999 2003 2009 Grass

Phylogeny Working Group 2001 Mabberley 2008) Reaching such a consensus is vital

since what we know of angiosperm phylogeny allows a very large number of

classifications to be based on it and as we find out more the number of possible

classifications increases greatly Unfortunately however nature does not dictate what

the classification should be All classifications are constructed by humans to

communicate particular aspects of groups and relationships Our goals as systematists

are surely to produce robust hypotheses of relationships to understand the evolution of

morphology and the like - but not to argue ad nauseam whether something should be

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

12

a family or a subfamily That way surely lies madness and worse the discredit of our

discipline

There are similar issues whatever naming system is used Thus in phylogenetic

naming (Baum et al 1998 for an example but cf Baum et al 2004 for the PhyloCode

see Cantino amp De Queiroz 2006) an unflagged hierarchy is used in which any

terminations of names used are uninformative about the relative position of taxa If one

adopts the principle of phylogenetic naming one indeed does not have to worry about

the nomenclatural consequences caused by lumping or splitting any well-supported

clade can be named without affecting the name of more or less inclusive clades

Unfortunately unflagged hierarchies have very serious deficiences as communication

devices because they lack one aspect essential in language biological or otherwise -

they contain no intrinsic information about the relationships of the group in question to

others (eg Pfeil amp Crisp 2005 Stevens 2006a) Recent suggestions for using prefixes like

Apo- and Pan- to PhyloCode names will however allow limited information of this

kind to be conveyed but only as it pertains to individual branches and current

proposals do not even mandate that the prefixes be employed consistently In any

event such proposals simply prevent the potential tripling of the number of quite

different names used to describe different aspects of a phylogenetic tree over those

used to name monophyletic groups pure and simple In general where n is the number

of extant species in a group the number of clades in such a group = n-1 (Species will

also need names too for their names see Dayrat et al 2008) Importantly here too

consensus over the clade names commonly learned by students and used in herbaria is

needed otherwise communication will be impeded the names themselves will provide

no guidelines as to which should be chosen The situation is of course more complicated

than this Terminations that convey ideas of rank in a phylogenetic classification can also

be used in phylocode names - however there they will carry no implications of rank

How they will be used is another matter of course

Of course there are other philosophies of classification and some still prefer

evolutionary classifications There classificatory principles differ substantially from those

followed here eg the recognition of paraphyletic taxa may be permitted however

detailed reasons for prefering the taxa that are recognised are rarely given although

nature and natural groups are often mentioned (cf Stuessy amp Koumlnig 2008) For

summaries of commonly used systems see Brummitt (1992) and Mabberley (2008)

new evolutionary systems appear every year or so However even those who allow or

promote the recognition of paraphyletic groups (eg Grant 2003 Thorne 2007

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

13

Heywood et al 2007) may find it of interest to examine a system recognizing only

monophyletic groups it provides a rather different understanding of evolution

Evolutionary classifications in general try and combine phylogeny and

morphological gaps although that is no easy thing to do - it is akin to combining chalk

and cheese (for an attempt to make this impossible task seem to be more objective see

Stuessy amp Koumlnig 2008) However there is a principle from evolutionary classification

that is relevant and I think quite useful here The size of the gap between two groups

tends to be inversely proportional to the sizes of the groups involved (Davis amp Heywood

1963) One can imagine a situation in which a large group is formally divided even

although the distinguishing characters of the two are weak whereas a smaller group

similarly divisable will be left intact

To summarize If hypotheses of phylogeny remain stable we should be able to base

a stable classification on that phylogeny and then get on with our work that is testing

the phylogenies we have elucidating phylogenies in areas where relationships are

unclear studying the evolution of morphology describing species etc In this context

the spread of the Angiosperm Phylogeny Group system (see below) and its widespread

utilisation in technical literature also floras (eg van der Meijden 2005) textbooks (eg

Simpson 2006 Judd et al 2007 [third edition]) dictionaries (Mabberley 2008) more

popular literature (eg Souza amp Lorenzi 2005 Spears 2006) and as an outline for a new

herbarium sequence (Haston et al 2007) is gratifying The posibility that one might be

able to develop a stable phylogeny-based classification of families and in particular

orders represents a dramatic turn-around from the pessimistic attitude about such

higher-level groupings expressed by Davis and Heywood (1963) and Thorne (1976) the

latter even suggesting that we should bury forever the metaphor of the phylogenetic

tree as highly unrealistic (ibid p 56) Returning to Godfray and Knapps (2004) users

of classifications who want a stable informative and accessible classification that

enables easy identification - unfortunately they want cake with everything and cannot

get it - these pages attempt to satisfy as many of their needs as possible but phylogeny

and monophyly are the primary shapers of the APG classification

On this classification in particular

Here I very largely follow the Angiosperm Phylogeny Group classification (APG

2003) Any differences are not to be interpreted as differences in principle simply that

new phylogenies continue to be published and here I attempt to provide an overview

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

14

of current ideas of higher-level relationships of seed plants The Angiosperm Phylogeny

Group classification is based on well-supported relationships evident in the numerous

molecular studies that began to appear in the late 1980s (see APG 1999 for the

principles underlying the classification) but recent phylogenetic work does not

contradict the major outlines of the trees used by APG II (2003) or even those in APG

I (APG 1999) The APG classification has for the most part been conservative and I

am similarly conservative here For the names and authorities I follow APG III (2009)

although the names of the authors may not always be correct these being in something

of a state of flux for the superordinal names used see Chase and Reveal (2009)

although classification at this level here is not exhaustive - only clades with several

orders have superordinal names

Some changes in our ideas of relationships and hence in the clades we talk about

are particularly likely in parts of Lamiales Caryophyllales and Malpighiales for example

although this will not affect the orders themselves Thus some changes to clade

circumscriptions are to be expected although I do not expect them to be substantial

but changes are neither a defect of cladistics nor a necessary consequence of the use of

molecular data Clades are hypotheses of relationships and as hypotheses they may be

overturned However work since APG I has not suggested other than minor changes

in the compositions of the orders even if the odd genus or even family is turning out to

be seriously misplaced - recent examples are Hydatellaceae (from monocots-Poales to

Nymphaeales Saarela et al 2007) Guamatelaceae (from Rosales-Rosaceae to

Crossosomatales Oh amp Potter 2006) and Perrottetia and Bhesa (from Celastrales-

Celastraceae to Huerteales and Malpighiales respectively Zhang amp Simmons 2006) The

main changes have been clarification of the relationhips of individual families or groups

of families that were of uncertain position eg of Chloranthaceae (Moore et al 2007)

Ceratophyllaceae (Jansen et al 2007) and unplaced asterid II families (Winkworth et al

2008a) As mentioned above phylogenies in the area of main interest in these pages are

overwhelmingly tree-like Although there is evidence for acquisition of host plant genes

by parasites (see below) even wider but inexplicable transfer of mitochochondrial

genes in plants like Amborella (Bergthorsson et al 2004 cf Goremykin et al 2009a)

and even transfers of nuclear genes (Vallenback et al 2008) these are the exception

rather than the rule

In cases where the Angiosperm Phylogeny Group II allowed alternatives as to the

limits of families - Papaveraceae in the broad sense or Papaveraceae plus

Pteridophyllaceae plus Fumariaceae Proteaceae in the broad sense or Proteaceae plus

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

15

Platanaceae - the choices made here follow common usage eg as in textbooks like

Judd et al (2007) and Simpson (2006) and particularly in the new edition of

Mabberleys The Plant Book (Mabberley 2008) This latter is itself an attempt to reflect a

consensus the result of taking the opinions of botanists at several meetings A largely

similar consensus is reflected in the APG III classification (APG 2009) where you will

see that alternative classifications have been dispensed with For many people the

existence of such alternative classifications will simply confuse so agreement over

which groupings to use when alternatives are permitted and then dispensing with the

alternatives that are not commonly used seems reasonable I am making the few

changes that are needed to bring the classification here in line with these others but

only gradually since there is not enough time in each day

As already mentioned higher-level relationships in general and the composition of

orders in particular have always presented something of a challenge to systematists

(eg Davis amp Heywood 1963) That being said the composition of some clades like

Apiales Crossosomatales and Pandanales is decidedly unexpected However it is

interesting to see that these higher level clades are generally accepted even in works

with different classificatory philosophies (eg Heywood et al 2007) For clades like

Malpighiales many of the family groupings within Asparagales etc attempts to find

distinctive characters have largely failed (but see Endress amp Matthews 2006a also

the Apomorphies page here) Interestingly as with families some groupings suggested

by molecular studies are supported by morphological andor chemical characters that

have long been known sometimes for over a hundred years the relationship between

Pittosporaceae and ApiaceaeAraliaceae are a case in point (Hegnauer 1969b and

references) As our knowledge of morphology and chemistry improves we can hope for

improvements in the characterisations of clades at all levels

In many cases the new family limits of the Angiosperm Phylogeny Group (see

APG 1999 2003) are not really controversial although changes from the limits

commonly accepted only a decade ago are sometimes quite dramatic (eg Wagenitz

1997) Thus the split of the old Saxifragaceae sl is necessitated by its extreme

polyphyly as also with Icacinaceaesl and Cornaceae sl However the limits and

relationships of such groups had long been considered to be unsatisfactory but until

recently there had been no real reason to prefer one proposed arrangement over

another It is generally accepted that the limits of Lamiaceae and Verbenaceae have to

be redrawn although there is as yet no compelling evidence that the redrawn taxa are

not sister taxa (there is no evidence that they are) But whatever their relationships the

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

16

content of the clades has changed considerably and incidentally they are now easier to

identify than before the decision to recognise two families is not difficult The same is

true for Salicaceae and Achariaceae (Malpighiales) two previously small families that

have received the bulk of the old Flacourtiaceae Clade and hence taxon limits remain

difficult around Theaceae (Ericalessee alsoSladeniaceae Pentaphylacaceae)

and Euphorbiaceae (Malpighiales see

also Phyllanthaceae Picrodendraceae Putranjivaceae etc) Here current groupings

may not represent quite such dramatic changes in our understanding of relationships

Indeed parts of the old Euphorbiaceae that were separated may yet go back together

particularly Phyllanthaceae and Picrodendraceae (see Wurdack et al 2004 Davis et al

2005) even if Theaceae should indeed be dismembered (Geuten et al 2004) However

given that there is no molecular evidence that currently warrants combining all the

segregates of Euphorbiaceae and even if some do come together the clades the

families represent suggest novel groupings not recognised in current classifications

Furthermore Rafflesiaceae appear to be embedded within Euphorbiaceae s str (Davis

et al 2007) so the family is divided Of course maintaining even a moderately narrowly-

circumscribed Euphorbiaceae would entail reducing the iconic Rafflesiaceae to

synonymy Relationships in core Caryophyllales especially

around Phytolaccaceae andNyctaginaceae and also Portulacaceae are incompletely

understood and refashioning of taxon limits will doubtless be needed as cladistic

relationships become apparent (eg see Nyffeler 2007 Ogburn 2007a b) Some

groupings in the old Icacinaceaeand Olacaceae also remain unclear

The discovery of the relationships of parasitic and aquatic groups have presented a

particular challenge to systematists Morphologically some of these plants are so highly

modified that interpretation of the plant body in conventional terms is difficult or even

impossible Thus parasitic groups (broad-leaved angiosperms only) such as Rafflesiaceae

are hard to place since both the vegetative body and the flowers are changed almost

beyond recognition (flowers of rafflesiaceae are a good example of gigantism - Davis et

al 2007) Furthermore plastid gene sequences may be difficult or impossible to obtain

the chloroplast DNA in particular being highly degraded and the rate of molecular

change in general may be high so the problem of long-branch attraction is serious (eg

Duff amp Nickrent 1997 Nickrent et al 1998 Caddick et al 2002a G Petersen et al

2006b) Echlorophyllous sapromycoheterotrophic taxa mostly known from monocots

present similar problems However progress is being made placements for

Apodanthaceae Rafflesiaceae Mitrastemonaceae Cytinaceae and Cynomoriaceae have

recently been suggested (Barkman et al 2004 2007 Davis amp Wurdack 2004 Nickrent et

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

17

al 2004 Davis et al 2007) it seems likely that Burmanniaceae sl are polyphyletic but

both parts are in Dioscoreales (Merckx et al 2006) and relationships within the largely

hemiparasitic Santalales are also gradually being clarified (Maleacutecot 2002) The inclusion

of parasitic taxa in general molecular analyses can cause conniptions (eg Nickrent et al

2004 Davis et al 2004 Chase et al 2006 G Petersen et al 2006b) and there can be

horizontal transmission of genes (eg Davis amp Wurdack 2005 Vitaceae to Rafflesiaceae

Barkman et al 2007 the mitochondrial atp1 gene commonly moves)

Similarly in water plants neither vessels in particular nor much xylem in general is

needed leaves are highly modified and water-mediated pollination if adopted may

well be associated with major changes in floral morphology Here too recent molecular

studies suggest that aquatic groups with hitherto problematic relationships may find

homes Thus Podostemaceae are close to Clusiaceae and Calophyllaceae and are sister

to Hypericaceae (Malpighiales Kita amp Kato 2001) Hydatellaceae which used to be in

Poales are part of Nymphaeales (Saarela et al 2007) and Hydrostachyaceae may be

close to Hydrangeaceae (Cornales Xiang et al 2002) or perhaps should be placed in

Lamiales note that in the first two cases in particular there are morphological and

chemical features that support such a move If Podostemaceae are indeed close to

Clusiaceae I look forward to seeing hypotheses to explain how the dramatic changes in

the vegetative body that have made Podostemaceae so problematic for generations of

systematists took place That conventional wisdom has trouble in understanding or

explaining how the morphologies of groups like Clusiaceae and Podostemaceae can be

related is largely a problem with conventional wisdom Ceratophyllaceae seem to be

finding a position as sister to eudicots (see Moore et al 2007) but their morphology is

so derived that there is no morphological evidence of which I am aware for this

relationship There are also similar although less extreme morphological problems with

many plants pollinated by wind

In such situations especially with some water plants mycoheterotrophs and

parasites a variety of characters may be affected and particularly under such

circumstances groups have tended to be formed using a few or even only a single

character that seems to provide evidence of relationships Add to this the tendency to

weight some characters particularly strongly on a priori grounds and the result was

either the recognition of conglomerate taxa such as Amentiferae which now appear to

be highly polyphyletic or the segregation of families like Plantaginaceae s str (now

much expanded) and Leitneriaceae (now in Simaroubaceae) - all of these are more or

less wind pollinated

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

18

Even although relationships of many parasitic and aquatic groups are being

clarified and at least some of the proposed phylogenetic positions seem to be well

supported it is indeed often still very difficult to understand how such groups have

evolved from their more morphologically conventional relatives However we have had

all sorts of preconceptions as to how characters should evolve Thus we used to

assume that features like highly scalariform vessel perforation plates or the complete

absence of vessels or a flower with an androecium that had many stamens a superior

ovary or separate petals were necessarily plesiomorphic or primitive and conversely

a simple perforation plate and the presence of vessels an androecium with few

stamens inferior ovary or petals that were connnate were almost necessarily

apomorphic or advanced (but cf eg Stebbins 1951) Such assumptions are incorrect

(eg Soltis et al 2005b) Carpels may become secondarily free carpels may fail to close

the seeds then developing outside the confines of the carpel as in some Aspagaraceae-

Nolinoideae Violaceae Berberidaceae Malvaceae-Sterculioideae etc) in Peliosanthes

teta perhaps the only species in Peliosanthes(Asparagaceae-Nolinoideae) the ovary

varies from superior to inferior (Jessop 1976 species limits here need close

investigation see also Kuzoff et al 2001 and Soltis amp Hufford 2002 Saxifragaceae

Apiales etc) many-seeded carpels can evolve from few-seeded carpels

(Razafimandimbison et al 2008) monoecy may be derived from dioecy (Schaefer amp

Renner 2010 and references) Classic studies such as those by Babcock (eg 1947)

on Crepis that assumed that evolution - in this case of the karyotype in particular - was

unidirectional have needed comprehensive re-evaluation (Enke amp Gemeinholzer 2008)

Most if not all characters have reversed andor evolved in parallel as is clear in the

discussion of ovary position here especially in Poales Saxifragales and Asparagales and

also in comments on such morphologically distinctive taxa (when compared with their

immediate relatives) such as Menyanthaceae and especially Pittosporaceae Parallel

evolution may occur even at the level of amino acid substitution as in the independent

acquisition of the phosphoenolpyruvate carboxylase (pepC) gene in C4 photosynthesis in

grasses (Christin et al 2007b see also Blaumlsing et al 2000)

Thinking of how such characters have been used in the past in classifications at the

generic level the results have been similar to those just discussed above for parasitic

plants and others Relying too much on animal pollination syndromes to mark generic

boundaries has all too often led to taxa that are highly unsatisfactory phylogenetically

(see eg Acanthaceae Bignoniaceae Campanulaceae Ericaceae Lamiaceae

Orchidaceae) and over-reliance on characters of fruit and seed (see eg Brassicaceae

and Apiaceae) has also led to unsatisfactory generic limits Again the more general

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

19

problem is the use of one or a very few characters that have been weighted a priori to

structure classifications (see also Garciacutea et al 2009) although it is perhaps not

surprising that such an approach sometimes results in the delineation of taxa that have

indeed turned out to be quite distinct (cf eg Burtt Astragalus versus Oxytropis)

Taxa recognized here are monophyletic yet many are polythetic at the

morphological level that is they lack unique features characterizing (defining) all and

only members of that family they can be recognised phenetically only by the unique

combinations of characters that they posess This is the result of evolution any

synapomorphy characterizing a taxon may be lost or modified beyond easy recognition

in some of its members or the synapomorphy may appear to be identical to a feature

that has evolved in parallel in a quite unrelated plant That plant groups are polythetic is

almost as much a feature of monophyletic taxa as the paraphyletic taxa common in

evolutionary classifications Some families now include substantial variation as

phenetically distinct derived groups are placed in their proper phylogenetic position -

examples are Ericaceae which include the erstwhile Empetraceae a wind-pollinated

group that is florally very different and the overwhelmingly large-flowered and animal-

pollinated Plantaginaceae which now include several derived small-flowered aquatic

and wind-pollinated groups that were previously placed in separate families

Thus some families as delimited here may not be easy to recognize However

remember that detecting relationships - use whatever characters you can even if they

are not obvious - and naming a plant - focus on easy-to-see characters that may not

reflect relationships - are quite different problems Taxa although natural may not be

readily recognizable indeed it was in exactly this context that Lamarck worked out the

basic principles of writing dichotomous keys in 1778 (see Scharf 2007 for keys and the

like) Of course Lamarcks idea of nature was very different from ours - he thought that

there was some kind of continuum of form on which living organisms were to be

situated with no real gaps anywhere - but this meant that his genera (for example)

might well not be sharply distinct from each other even if each were part of the real

continuum that was life Lamarcks separation of the establishment of relationships and

groupings in a system and identification of the plants in that system have been widely

accepted (but cf eg Godfray amp Knapp 2004 many of the contributors to Heywood et

al 2007) Perhaps the best way of identifying plants at the family level is by well-made

multiple access keys as in Watson and Dallwitz (1992a onwards family limits there may

differ substantially from those adopted here) Multiple access keys free users from the

constraints of dichotomous keys in which particular characters are needed at each step

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

20

of the identification process before there can be further progress Instead those

characters that happen to be evident on a specimen can be used in whatever order is

convenient when linked to illustrations glossaries etc their power is enormous (see

Dallwitz et al 2000 [2006] for the principles underlying their construction and use)

Nevertheless dichotomous keys such as those of Hutchinson (1973) and Franz Thonner

(Geesink et al 1981) have their uses Of course taxa coming out adjacent in keys may

well not be at all related

(When identifying large numbers of plants even more efficient than either style of

identification and certainly lots more fun is sight identification Unless you have a

photographic memory you have to build up your knowledge of comparative plant

morphology - on which the ability to make accurate identifications depends - by

repeated observation When faced with an unknown plant I always look for leaf teeth

and stipules distinctive hair types especially stellate and T-shaped hairs glands and

punctations of any sort the presence of latex or other exudate and check leaf insertion

smelling crushed leaves can also be helpful In this context nodal anatomy can usually

be checked using a razor and a hand lens or even simply carefully examining leaf scars

The short paragraphs added after most families may help in confirming familial

identifications)

For the record and for the little that it is worth there are 4 orders and 13 families

of gymnosperms characterised on these pages and together they include some 82

genera and 947 species For angiosperms comparable figures are 56 orders 445

families 13208 genera and 261750 species (of which monocots include 11 orders 89

families 2759 genera and 52760 species) Note however that higher mathematics

was never my strong point and anyway these are pretty meaningless figures even for

species which many (but not all) might concede smacked slightly more of reality than

other taxa estimates range as high as 422000 (Govaerts 2001) Furthermore numbers

of genera and species change daily Nevertheless as emphasized here families are

useful in teaching we as a community can ensure that their limits remain largely stable

and by concentrating on relatively few of them one can gain some familiarity with much

of the worlds flora For summaries of the sizes of orders and families and of the general

arrangement followed in these pages see also the Statistics page itself

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

21

A PHYLOGENETIC TREE OF SEED PLANTS (THE MAIN TREE)

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

22

The Main Tree above is a conservative summary of well supported relationships

between the major clades of extant seed plants Reading - interpreting and

understanding - such a phylogenetic tree is not necessarily easy so reading a tree may

require a bit of practice OHara (eg 1988 1992) provides an accessible introduction to

thinking and talking about history in the context of looking at phylogenetic trees More

recently Krell and Cranston (2004) Crisp and Cook (2005) and others have emphasized

how careful one must be when interpreting and talking about evolution in ladderized

trees in particular and phylogenetic trees in general Gregory (2008) provides a

particularly detailed discussion of this problem In ladderized trees the smaller (in terms

of numbers of terminals) sister taxon is consistently shown on the same side at every

node thus the trees tend to be pectinate like the teeth of a comb however the

horizontal axis that is as a result so evident has no polarity or particular direction The

use of the adjective basal when discussing phylogenetic trees is especially dangerous

(see also Wojciechowski et al 2004 D Soltis et al 2005b) When I use the term and the

context is not otherwise clear I am referring to the pectinations at the base of a

ladderised tree Remember that when talking about sister taxa one can never be basal

to the other or older or younger although members of one can be more derived - in the

sense of having more apomorphies - than the other (but even then one has to be

careful) The words primitive and advanced and lower and higher should

practically never be used especially when talking about taxa Amborellaceae and

Pinaceae are sister to all other flowering plants and Pinales and are basal to the crown

groups of all other flowering plantsPinales respectively but that does not mean they

are thereby primitive Pinaceae in particular have numerous apomorphies The word

plesiomorphic is far less loaded than primitive and can be used to talk about

individual characters

Most trees here have been more or less ladderized as is the one above The

ladderization may be imperfect for example you can see that on the Main Tree

asterids with ten orders follow rosids and relatives which have seventeen although

this is in part because of the currently unresolved nature of relationships between the

main clades of core eudicots (this is likely to change soon) The Main Tree could have

been drawn with Amborella Acorus or a host of other taxa at the far right without

offending any relationships phylogenetic trees are like mobiles the only fixed points

being the nodes Despite the problems just mentioned pectination interpreted

carefully has its value and it is not an arbitrary process As one reads the terminals of a

pectinate tree from left to right adjacent terminals will be separated by apomorphies

that will cumulate in an appropriate fashion When thinking of a book or a herbarium

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

23

sequence (see Haston et al 2007) this is of some value Nymphaeales and

Austrobaileyales are here adjacent on the tree but they could be separated by

hundreds of families in the sequence merely by rotating only the node from which

Nymphaeales and their sister group (all other angiosperms minus about 75 species)

arise If adjacent in a book or herbarium or as in these summary pages then it is

relatively easy to relate their characters both apomorphic and plesiomorphic but if

separated by hundreds of pages or two floors in a large building then it is less easy to

get anything from the sequence (Since all orders in the pages of the main part of this

site are preceded by the apomorphies of all nodes immediately below them in the seed

plant phylogeny and because of the linkages that have been built in to the site this

problem is much less serious there there is no sequence) So the particular sequence of

all seed plants that is based on a phylogenetic classification but is to be used in herbaria

or any other place where taxa have to be arranged linearly as in this part of APweb

will be one that maximizes the number of taxa that are both successive branches of the

tree and placed successively in the sequence Since specimens are generally filed under

families the outline of a new family sequence for arranging herbaria and books can now

be suggested (Haston et al 2007) and such a standardized sequence will further help

teaching and learning about plants

ON THINKING ABOUT APOMORPHIES

Identifying apomorphies is important because understanding the evolution of

morphology in the broad sense - ie including anatomy chemistry etc - in the context

of diversification is one of our major goals For identifying apomorphies several

preconditions must be met One needs to have an accurate well supported phylogeny

one has to have examined the right taxa both from the point of view of morphology and

molecules one has to have coded the characters right (ie delimited states

appropriately) and one has to use an appropriate model of evolution when fitting the

variation to the tree (see eg Omland 1999 Stevens 2006b) If the distributions of

apomorphies below are compared with those in other studies (eg Turgeon et al 2001

Bremer et al 2001 Endress 2001 Albach et al 2001a Judd amp Olmstead 2004 D Soltis

et al 2005b Zhang et al 2006 Judd et al 2007) the reader may find a number of

differences Although I have integrated such studies as far as is possible there are five

reasons that link to the preconditions just mentioned why there may be differences

where features are placed on trees

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

24

1 I may not have found all the information about a particular character there may be

disagreement over its interpretation or I have added information

2 The sampling of nearly all molecular studies is very incomplete (see Salisbury amp Kim

2001 for problems caused by sampling) indeed it is commonplace to decry the

incompleteness of molecular sampling But not only is the sampling in molecular studies

often less than we might wish that of the morphological and chemical characters whose

evolution we are interested in understanding is also often very poor So for many

anatomical chemical and embryological characters that are confidently said to

characterise families and other groups we all too often have no idea if those characters

are applicable to the whole clade or just to a subgroup within that clade However as

our sampling improves we can locate changes on the tree more precisely if in

unexpected positions Thus is is sometimes suggested that Ericaceae have ellagic acid

(Soltis et al 2005b) in fact the little evidence we have indicates that only a small clade

of some 80 species has ellagic acid and the rest of the family - itself alone about one

third of all Ericales - does not Similarly Ericaceae are noted for their

ectendomycorrhizal associations but Enkianthus sister to all other Ericaceae

apparently lacks these (Abe 2005) Thus these mycorrhizae and their associated

distinctive hair roots are unlikely to be apomorphies of Ericaceae - however it appears

that other families near Ericaceae may also have ectendomycorrhizae (see Asai 1934)

3 I am fitting characters to a very conservative tree with many polytomies although the

nodes that are utilised are for the most part strongly supported polytomies make the

optimisation of characters that is the assigment of character state change to a

particular node on the tree notably difficult (eg Madison amp Madison 2002) In nearly

all studies of the evolution of characters distributions of characters are optimised on a

more or less fully resolved tree and the construction of supertrees may yield yet more

detailed hypotheses of relationships (for literature on supertrees see Cotton amp

Wilkinson 2007 2008) Of course some nodes on such fully resolved trees andor

supertrees may have little support and optimisations of characters on such trees may

carry correspondingly little conviction Even parts of some of the trees used here have

poor support eg relationships within aquatic Alismatales etc although I indicate

when this is the case

4 Exactly how one goes about optimising a character on a tree is critically important

Even using simple parsimony optimisations (ACCTRAN or DELTRAN ACCelerated

TRANsitions or DELayed TRANsitions) the position of synapomorphies on trees - and

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

25

hence our ideas of evolution - may differ and this is still more true if one uses maximum

likelihood or Bayesian methods and makes apparently reasonable suggestions about

weighting gains over losses (eg Donoghue amp Ackerley 1996 Cunningham et al 1998

Omland 1997 1999 Ree amp Donoghue 1999 Polly 2001 Webster amp Purvis 2001

Ronquist 2004 Crisp amp Cook 2005) Sannier et al (2007) show how in in Arecaceae that

where on a phylogenetic tree one might peg changes in microsporogenesis will depend

on the methods one uses to do this and Pedersen et al (2007) discuss the sometimes

very substantial effect of node support on the posterior probabilities of ancestral

character states Here I use parsimony optimization not always as explicit as it might be

but I have often indicated where there are particularly important uncertainties as to the

positions of particular character changes on the tree

5 Finally although I have paid quite a lot of attention to the delimitation of the

character states that make up all the characterizations I have not spent enough time on

this critical operation If we are interested in understanding evolution then fitting the

basic variation - not character states - to a tree in principle allows greater flexibility in

understanding morphology in the context of local phylogenies (see also Stevens 2000

Endress 2005c) However many character states used here are delimited globally that

is they are circumscribed in the context of the variation shown by individual characters

across all angiosperms andor in the context of classic ideas of character evolution

Character states often have arbitrary limits and serve best to communicate

information whether they are in fact suitable for either phylogenetic analysis or

understanding evolution are separate issues Studies have rather unsurprisingly

perhaps but importantly shown that dividing the one character into different sets of

states may yield differing ideas of evolution of that character (eg Lamb Frye amp Kron

2003 Hibbett 2004) When looking at trees on which character states are optimised

one should bear in mind the problems surrounding the delimitation of states (eg

Stevens 2000 2006b) and the danger of using pollen or other types - constructs

based on many characters that vary independently but which effectively get lost in

these types thus Blackmore et al (2009) decomposed the pollen types in Asteraceae

into 52 characters

I use Remanes three main criteria of homology or better similarity when

determining the basic similarity of structures on different organisms (see Remane 1952)

These criteria are special properties position and intermediates Special properties

include anatomical or chemical characters gene expression data etc Position refers

to the position of an organ with respect to landmarks on the plant Although plants are

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

26

plants and landmarks are not as fixed as one might like trying to work out the relative

positions of parts is a good way of understanding morphology so long as one does not

try to out-do Procrustes Intermediates include intermediates found on the same

plant or on different plants Intermediates may observed during development or be

adult structures Thus one may find changes during development which suggest what

the nature of a particular structure is and very different stuctures on different plants

may turn out to be similar early in development Or one can simply compare different

taxa and see that structures that appear to be very different when comparing two

different taxa can be linked morphologically by looking at other taxa

Nevertheless the use of these criteria may not yield an unambiguous answer as to

what a structure is even given a solid phylogeny and an improved understanding of

development (see Jaramillo amp Kramer 2007 for a useful discussion) As Endress (2005c)

observed a number of features - position function development shape anatomy

histology gene activity and relationships to other taxa that clearly have petals - can be

used to distinguish a petal (for example) from other floral structures if a petal does not

have one of these features is it thereby not a petal Thus Maturen et al (2005) recently

found that floral organ diversity genes (B and C) were expressed in the large white

inflorescence bracts of Cornus (see also Costa et al 2005) Peney et al (2005) noted that

not all monosulcate pollen grains in monocots have the same developmental pathway

and that as a result such pollen might not have the same ancestral state Reeves and

Olmstead (2003) suggested that the genetic mechanisms causing monosymmetry in

Lamiales and Solanales were different and Serna and Martin (2006) described similar

problems with the development of hairs in Arabidopsis when compared with that of

hairs in Antirrhinum and Solanaceae Indeed as one perhaps might expect delimitation

of states and characters does not necessarily become easier with increasing knowledge

of development etc Thus Buzgo et al (2004) Matthews and Endress (2005) and others

have shown how hard it can be to distinguish between eg prophylls and other floral

structures as their behaviour is studied during the course of the development of the

flower Similarly at what concentration is a particular secondary metabolite deemed to

be present (Waterman (2007)

The validity of the approach used here that of fitting morphological variation to a

largely molecular-based tree may be questioned However I think it rather unlikely that

well-supported molecular branches will be overturned by morphological data Indeed

analyses of morphological data alone do provide support for many of the clades evident

in molecule-only analyses and analyses with morphological and molecular data

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

27

together may lead to increased support for clades (eg Hufford 1992 Nandi et al 1998

[but adding morphological data reduces support for a number of critical clades too]

Doyle amp Endress 2000 however in none of these papers is the use of morphology

without ambiguity) It is unfortunately clear that the use of morphology alone may not

suggest problems in the phylogenetic placement of taxa that later turn out to have been

wrongly included (eg Zhang et al 1992) Thus I have been wary of putting much weight

on clades that have only morphological support but note that I have treated molecule-

based clades with low bootstrap or jacknife support values (esp below 70) or low

posterior probabilies (below 095) likewise Although I may have been mistaken in

placing so much emphasis on molecular data in terms of providing the basic

phylogenetic framework for angiosperms morphological and molecular data are only

very rarely in irreconcilably strong conflict There are indeed a few places where the

conflict seems extreme These include the relative positions of the Monimiaceae and

Hernandiaceae (Laurales) the position of Hanguanaceae (Commelinales [as here] or

Zingiberales) and of Triplostegia (is it in Dipsacaceae or Valerianaceae - see

Dipsacales) Fossils are unlikely to affect the topologies of the trees presented here but

see below for their importance in understanding morphological evolution in general and

the evolution of angiosperms in particular Nevertheless some largely reject the idea

that trees based on molecular data alone can recover phylogenetic relationships

especially when branching points are old and prefer to used trees based on analysis of

morphological data including those taken from fossils (Hilton amp Bateman 2006 Farjon

2007)

All in all however the extent of the congruence between morphological and

molecular data is impressive and heartening and many clades can be characterised

morphologically It seemed in 1998 that there were no unambiguous morphological

synapomorphies for angiosperm orders (K Bremer 2000) and this is still true if by

unambiguous is meant non-homoplasious However many orders and other clades

have synapomorphies even if these may be indistinguishable at least at the current

level of morphological and developmental knowledge from parallel occurences

elsewhere As our morphological knowledge increases so too does the number of

apomorphies

Endress and Matthews (2006a) emphasize the importance of tendencies and

developmental constraints when thinking about characters of clades within the rosids -

many characters or character combinations occur in a rather sporadic fashion within

the clade and apparently notably less frequently outside it There are a number of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

28

examples of tendencies here and in the individual order pages such as the features

enclosed by parentheses in groups above the level of family eg see (cuticular waxes

as aggregated rodlets) for the commelinids and the discussion of the distribution of

polyandry that is flowers with numerous stamens in the asterid I + II groups etc

Tendencies may involve more than single characters When thinking of the

associations of features like integument thickness and vascularization of the integument

(Endress amp Matthews 2006a) one can imagine fairly simple developmental

preconditions being operative As Endress and Matthews (2006a) note it is difficult to

imagine vascular tissue developing in an integument only two cells thick Changes here

would be loosely correlated if morphologically linked However the strongly correlated

changes noted by Givnish et al (2005) are ecologically linked but are presumably

morphologicallydevelopmentally independent When there seem to be characters

evolving more or less together tests can be carried out to see if the changes are

concentrated on certain branches of the tree (eg Maddison 1990 Sanderson 1991

Maddison amp Maddison 2000) Such tests have rarely been carried out at higher levels in

angiosperms

One of the most striking examples of a tendency is the distribution of N-fixation

restricted as it is to a monophyletic group of four clades although it has arisen seven (or

perhaps several more) times independently within the N-fixing clade and several

members of two quite different kinds of bacteria are involved (eg Clawson et al 2004

Elliott et al 2007 Sprent amp James 2007) Other examples of work that bears on the issue

of tendencies include the findings that flowers of polysymmetrical Arabidopsis have

genes like TCP1 that are expressed asymmetrically during early development and TCP1

is a probable orthologue of the well-known CYC gene of Antirrhinum that is involved in

the development of monosymmetric slowers there (Cubas et al 2001 Costa et al 2005

etc) Parallelism might build on this underlying morphologically cryptic monosymmetry

even if details of the genetic mechanisms causing the monosymmetry evident in

particular groups may be different (Reeves amp Olmstead 2003 Cubas 2004 see above)

Similarly there have been several recent suggestions that the capability to synthesise a

particular metabolite may be switched off but not lost and so can sometimes be

reacquired (eg Wink amp Witte 1983 Wink 2003 Liscombe et al 2005 Larsson 2007

Waterman 2007) Hence perhaps the rather spotty distribution of many secondary

metabolites like ellagic acid the indole alkaloid camptothecin iridoids etc when

considered in the context of phylogenies As a non-botanical example - but a rather nice

one - Salwini-Plawen and Mayr (1961) suggested some time ago that there has been

considerable parallelism (40-65 or more independent origins) in the evolution of eyes in

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

29

metazoans However Pax 6 seems to be a master control gene that is involved in eye

formation perhaps throughout all bilateralians (eg Gehring amp Ikeo 1999 Erwin amp

Davidson 2002) which changes the nature of the problem Note however that in

general our invocation of evolutionary tendencies to explain the patterns of

distributions of characters is really an admission that we do not understand those

patterns

As we find out more about variation we will probably find fewer and fewer features

constant throughout a group Most unqualified statements of presence and absence

should properly be qualified as usually present or usually absent if one is thinking of

the characterisations on the order pages as encompassing the total variation within a

clade Thus Pistia alone among monocots as so far known has sieve tube plastids with

starch grains not protein crystals However this does not affect the fact that sieve tube

plastids with cuneate protein crystals and lacking starch grains are an apomorphy for

monocots Furthermore fossils in a number of cases suggest character combinations

unknown in extant taxa as may be seen in the discussions of Fagaceae Platanaceae

Iteaceae Calycanthaceae etc Confusing the issue there may be questions as to where

exactly on the tree a particular fossil is to be placed (eg see Nymphaeaceae

Calycanthaceae Archaefructus etc)

To summarize given our current understandings of both phylogenies and

characters evolution of some characters in which we are interested seems very labile

(see eg D Soltis et al 2005b Endress and Matthews 2006a Stevens 2006b Ekman et

al 2008 for a good recent study) and I have been cautious when talking about character

evolution Much effort must continue to be spent in summarizing characters of well-

established clades at all levels providing features by which they may be recognized and

signaling synapomorphies Remember that (1) the basic morphological anatomical and

chemical knowledge of many critical taxa is woefully incomplete (2) different

assumptions about character evolution may greatly affect the position of

synapomorphies on trees (3) in many cases relationships within and between many

groups are too uncertain at present to worry very much about synapomorphies and

(4) we must be clear about what we do and do not not know As mentioned above I

have indicated in a number of places where there are particularly important

uncertainties as to where characters should be placed on the tree Nevertheless it is a

relatively easy matter to update notes such as these and it can be a simple matter to

incorporate new data on characters that have never before been considered in the

context of a tree Much basic - and unfortunately perhaps unfashionable - work must be

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

30

carried to clarify the distribution of morphological anatomical and chemical characters

for examples of what can to be done see the work of P K Endress and collaborators

(floral morphology and development) and S R Jensen and collaborators (iridoids) But

acquiring information about nodal anatomy cell and tissue distribution nodal anatomy

and the like is not difficult at all and should be generally encouraged

But all this is merely a necesary prelude to the understanding of evolution For this

one needs to know a lot more including dating the phylogeny understanding the

function(s) of characters factoring in relevant aspects of the palaeoenvironment in

which an apomorphy first appeared etc

SUMMARY OF THE SYSTEM

Below is a formalised summary of the relationships within orders of the families of seed

plants There are a few families that are not recognised even as options in APG II and

vice versa as well as a few extra orders The families to be recognised in the most

recent edition of Mabberleys The Plant Book (Mabberley 2008) are also largely

consistent with those below But all differences are trivial and will - I hope - eventually

disappear

Square brackets - [] - enclose clades the plus sign - + - designates sister taxa a

comma - - denotes part of a polytomy and quotation marks - - denotes a

paraphyletic group

SEED PLANTS

GYMNOSPERMS

Cycadales

Cycadaceae + Zamiaceae

Ginkgoales

Ginkgoaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

31

Pinales

Pinaceae [[Araucariaceae [Podocarpaceae + Phyllocladaceae]] [Sciadopityaceae

[Taxaceae + Cupressaceae]]]

Gnetales

Ephedraceae [Gnetaceae + Welwitschiaceae]

ANGIOSPERMSFLOWERING PLANTS

Amborellales

Amborellaceae

Nymphaeales

Hydatellaceae [Cabombaceae + Nymphaeaceae]

Austrobaileyales

Austrobaileyaceae [Schisandraceae + Trimeniaceae]

Chloranthales

Chloranthaceae

MAGNOLIIDS

Magnoliales

Myristicaceae [Magnoliaceae [[Himantandraceae + Degeneriaceae] [Eupomatiaceae +

Annonaceae]]]

Laurales

Calycanthaceae [[Siparunaceae [Gomortegaceae + Atherospermataceae]] [Monimiaceae

[Hernandiaceae + Lauraceae]]]

Canellales

Canellaceae + Winteraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

32

Piperales

[Hydnoraceae + Aristolochiaceae] [Piperaceae + Saururaceae]

MONOCOTS

Acorales

Acoraceae

Alismatales

Araceae [Tofieldiaceae [[Alismataceae [Hydrocharitaceae + Butomaceae]]

[Scheuchzeriaceae [Aponogetonaceae [Juncaginaceae [Maundiaceae [[Posidoniaceae

[Ruppiaceae + Cymodoceaceae]] [Zosteraceae + Potamogetonaceae]]]]]]]

Petrosaviales

Petrosaviaceae

Dioscoreales

Nartheciaceae [[Taccaceae + Thismiaceae] [Burmanniaceae + Dioscoreaceae]]

Pandanales

Velloziaceae Triuridaceae Stemonaceae [Pandanaceae + Cyclanthaceae]

Liliales

Corsiaceae [Campynemataceae [Petermanniaceae [Colchicaceae + Alstroemeriaceae]]

Melanthiaceae [[Philesiaceae + Rhipogonaceae] [Smilacaceae + Liliaceae]]]

Asparagales

Orchidaceae [[Boryaceae [Blandfordiaceae [Lanariaceae [Asteliaceae + Hypoxidaceae]]]]

[[Ixioliriaceae + Tecophilaeaceae] [Doryanthaceae [Iridaceae [Xeronemataceae

[Xanthorrhoeaceae [Alliaceae + Asparagaceae]]]]]]]

COMMELINIDS

Unplaced

Dasypogonaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

33

Arecales

Arecaceae

Poales

[Typhaceae + Bromeliaceae] [Rapateaceae [[Thurniaceae [Juncaceae + Cyperaceae]]

[[Mayacaceae [Eriocaulaceae + Xyridaceae]] [Flagellariaceae [Anarthriaceae

[Centrolepidaceae + Restionaceae]]] [Joinvilleaceae [Ecdeiocoleaceae + Poaceae]]]]]]]

Commelinales

[Commelinaceae + Hanguanaceae] [Philydraceae [Haemodoraceae + Pontederiaceae]]

Zingiberales

Musaceae [Strelitziaceae + Lowiaceae] Heliconiaceae [[Cannaceae + Marantaceae]

[Costaceae + Zingiberaceae]]

Ceratophyllales

Ceratophyllaceae

EUDICOTS

Ranunculales

Eupteleaceae [Papaveraceae [[[Lardizabalaceae + Circaeasteraceae] [Menispermaceae

[Berberidaceae + Ranunculaceae]]]]

Sabiales

Sabiaceae

Proteales

Nelumbonaceae [Platanaceae + Proteaceae]

Trochodendrales

Trochodendraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

34

Buxales

Haptanthaceae [Buxaceae + Didymelaceae]

CORE EUDICOTS

Gunnerales

Gunneraceae + Myrothamnaceae

Dilleniales

Dilleniaceae

Saxifragales

Peridiscaceae [[Paeoniaceae [Altingiaceae [Hamamelidaceae [Cercidiphyllaceae +

Daphniphyllaceae]]]] [[Crassulaceae [Aphanopetalaceae [Tetracarpaeaceae

[Penthoraceae + Haloragaceae]]]] [Iteaceae [Grossulariaceae + Saxifragaceae]]]]

Cynomoriaceae unplaced

Vitales

Vitaceae

ROSIDS

FABIDROSID I

Zygophyllales

Krameriaceae + Zygophyllaceae

Celastrales

Lepidobotryaceae + Celastraceae

Oxalidales

Huaceae [[Connaraceae + Oxalidaceae] [Cunoniaceae [Elaeocarpaceae [Brunelliaceae +

Cephalotaceae]]]]

Malpighiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

35

[Achariaceae [Goupiaceae [Violaceae + Passifloraceae] [Lacistemataceae Salicaceae]]

[Lophopyxidaceae Putranjivaceae] [Ctenolophonaceae [Erythroxylaceae +

Rhizophoraceae]] Linaceae Ixonanthaceae Humiriaceae Irvingiaceae

Centroplacaceae Pandaceae [[Bonnetiaceae + Clusiaceae] [Calophyllaceae

[Hypericaceae + Podostemaceae]]] [Malpighiaceae + Elatinaceae] Ochnaceae

[Peraceae [Rafflesiaceae + Euphorbiaceae]] [Phyllanthaceae + Picrodendraceae]

[Balanopaceae [[Trigoniaceae + Dichapetalaceae] [Chrysobalanaceae +

Euphroniaceae]]] Caryocaraceae

N-FIXING CLADE

Fabales

Quillajaceae [Fabaceae [Polygalaceae + Surianaceae]]

Rosales

Rosaceae [[Barbeyaceae [Elaeagnaceae Dirachmaceae Rhamnaceae]] [Ulmaceae

[Cannabaceae [Moraceae + Urticaceae]]]]

Cucurbitales

Anisophylleaceae [[Corynocarpaceae + Coriariaceae] [Cucurbitaceae [Tetramelaceae

[Datiscaceae + Begoniaceae]]]] Apodanthaceae

Fagales

Nothofagaceae [Fagaceae [[Myricaceae + Juglandaceae] [Casuarinaceae

[Ticodendraceae + Betulaceae]]]]

MALVIDROSID II

pgtGeraniales

Geraniaceae [[Melianthaceae + Francoaceae] [Vivianaceae + Ledocarpaceae]]

Myrtales

Combretaceae [[Onagraceae + Lythraceae] [[Vochysiaceae + Myrtaceae]

[Melastomataceae [Crypteroniaceae [Alzateaceae + Penaeaceae]]]]]

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

36

Crossosomatales

[Staphyleaceae [Guamatelaceae [Crossosomataceae + Stachyuraceae]]] [Aphloiaceae

[Geissolomataceae + Strasburgeriaceae]]

Picramniales

Picramniaceae

Sapindales

Biebersteiniaceae Nitrariaceae [Kirkiaceae [Anacardiaceae + Burseraceae]]

[Sapindaceae [Simaroubaceae Rutaceae Meliaceae]]

Huerteales

Gerradinaceae [Dipentodontaceae + Tapisciaceae]

Malvales

Neuradaceae [Thymelaeaceae [Sphaerosepalaceae Bixaceae [Cistaceae

[Sarcolaenaceae + Dipterocarpaceae]] [Cytinaceae + Muntingiaceae] Malvaceae]]

Brassicales

[Akaniaceae + Tropaeolaceae] [[Moringaceae + Caricaceae] [Setchellanthaceae

[Limnanthaceae [[Koeberliniaceae [Bataceae + Salvadoraceae]] [Emblingiaceae

[Pentadiplandraceae [Gyrostemonaceae Resedaceae Stixis and relatives] Tovariaceae

[Capparaceae [Cleomaceae + Brassicaceae]]]]]]]]

Berberidopsidales

Aextoxicaceae + Berberidopsidaceae

Santalales

Erythropalaceae Olacaceae [[Loranthaceae [Misodendraceae + Schoepfiaceae]]

[Opiliaceae + Santalaceae]] Balanophoraceae unplaced

Caryophyllales

[[Droseraceae [Nepenthaceae [Drosophyllaceae [Ancistrocladaceae +

Dioncophylleaceae]]]]] [[Frankeniaceae + Tamaricaceae] [Polygonaceae +

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

37

Plumbaginaceae]]] [Rhabdodendraceae [Simmondsiaceae [[Asteropeiaceae +

Physenaceae] [[Caryophyllaceae [Achatocarpaceae + Amaranthaceae]]

[Stegnospermataceae [Limeaceae [[Lophocarpaceae [Barbeuiaceae [Aizoaceae

[Sarcobataceae Phytolaccaceae Nyctaginaceae]]]] [Molluginaceae [Halophytaceae

Montiaceae Basellaceae Didiereaceae [Talinaceae [Portulacaceae [Anacampseros etc +

Cactaceae]]]]]]]]]]]]]

ASTERIDS

Cornales

[Cornaceae + Nyssaceae] [Hydrangeaceae + Loasaceae] Hydrostachyaceae [Curtisiaceae

+ Grubbiaceae]

Ericales

[Balsaminaceae [Marcgraviaceae + Tetrameristaceae]] [[Polemoniaceae +

Fouquieraceae] Lecythidaceae [[Sladeniaceae + Pentaphylacaceae] [Sapotaceae

[Ebenaceae [Maesaceae [Theophrastaceae [Myrsinaceae + Primulaceae]]]]]

[Mitrastemonaceae Theaceae [Symplocaceae [Styracaceae + Diapensiaceae]]

[[Sarraceniaceae [Roridulaceae + Actinidiaceae] [Clethraceae [Cyrillaceae +

Ericaceae]]]]]]

LAMIIDASTERID I

Unplaced

Metteniusaceae Oncothecaceae Icacinaceae and some unplaced ex-Icacinaceae

Garryales

Garryaceae + Eucommiaceae

Unplaced

Boraginaceae Vahliaceae

Gentianales

Rubiaceae [Gentianaceae [Loganiaceae [Gelsemiaceae + Apocynaceae]]]

Lamiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

38

Plocospermataceae [Oleaceae [Tetrachondraceae [Calceolariaceae

[Gesneriaceae Peltanthera [Pedaliaceae Carlemanniaceae Martyniaceae Stilbaceae

Plantaginaceae Linderniaceae Bignoniaceae Verbenaceae Lamiaceae Paulowniaceae

Schlegeliaceae Thomandersiaceae Phrymaceae Rehmannia Acanthaceae

Scrophulariaceae Orobanchaceae Byblidaceae Lentibulariaceae]]]]]

Solanales

[Montiniaceae [Sphenocleaceae + Hydroleaceae]] [Convolvulaceae + Solanaceae]

CAMPANULIDASTERID II

Aquifoliales

[[Cardiopteridaceae + Stemonuraceae] [Aquifoliaceae [Helwingiaceae +

Phyllonomaceae]]

Asterales

[Rousseaceae + Campanulaceae] Pentaphragmataceae [[Alseuosmiaceae [Phellinaceae

+ Argophyllaceae]] Stylidiaceae [Menyanthaceae [Goodeniaceae [Calyceraceae +

Asteraceae]]]]

Escalloniales

Escalloniaceae

Bruniales

[Bruniaceae + Columelliaceae]

Apiales

Pennantiaceae [Torricelliaceae [Griseliniaceae [Pittosporaceae [Araliaceae

[Myodocarpaceae + Apiaceae]]]]]

Paracryphiales

Paracryphiaceae

Dipsacales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

39

Adoxaceae [Diervillaceae [Caprifoliaceae [Linnaeaceae [Morinaceae [Dipsacaceae +

Valerianaceae]]]]]

LINKS TO ORDERS AND FAMILIES

[Back to Top]

MAIN GROUPINGS

Asterids commelinids core eudicots asterid 1 asterid

2 eudicots gymnosperms Magnoliophyta monocots N-fixing claderosids seed plants

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A | B | C | D | E | F | G | H | I | J | L | M | N | O | P | Q | R | S | T | U | V | W | X | Z

Abietales Acanthales Acerales Acorales Actinidiales Actinostrobales Adoxales Aescul

ales Agavales Aizoales AkanialesAlismatales Alliales Alseuosmiales Alstroemeriales

Altingiales Amaranthales Amaryllidales Amborellales AmbrosialesAmmiales Amomal

es Ancistrocladales Anisophylleales Annonales Anthobolales Apiales Apocynales Apo

nogetonalesAquifoliales Arales Araliales Aralidiales Araucariales Arecales Aristolochi

ales Athrotaxidales Asarales AsclepiadalesAsparagales Asphodelales Asparagales Ast

eliales Atriplicales Aucubales Austrobaileyales Avenales

Balanitales Balanopales Balanophorales Balsaminales Barbeyales Barclayales Batales

Begoniales BerberidalesBerberidopsidales Betulales Biebersteiniales Bignoniales Bix

ales Boraginales Brassicales Brexiales Bromeliales BrunialesBrunoniales Burmannial

es Burserales Butomales Buxales Byblidales

Cactales Callitrichales Calycanthales Calycerales Campanulales Campynematales Can

ellales Cannales CapparalesCaprifoliales Cardiopteridales Carduales Caricales Carle

manniales Caryophyllales Cassiales Casuarinales CelastralesCentrolepidales Cephalot

ales Cephalotaxales Ceratophyllales Cercidiphyllales Chenopodiales Chironiales Chlor

anthalesChrysobalanales Cinchonales Circaeasterales Cistales Citrales Cocosales Col

chicales Columelliales CombretalesCommelinales Connarales Convolvulales Coriarial

es Cornales Corylales Corynocarpales Crassulales CrossosomatalesCucurbitales Cunn

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

40

inghamiales Cunoniales Cupressales Cycadales Cyclanthales Cymodoceales Cynarales

CynomorialesCyperales Cyrillales Cytinales

Daphnales Daphniphyllales Dasypogonales Datiscales Degeneriales Desfontainiales D

iapensiales Didymelales DillenialesDioales Dioncophyllales Dioscoreales Diospyrales

Dipentodontales Dipsacales Droserales

Ebenales Echiales Elaeagnales Elaeocarpales Elatinales Elodeales Emmotales Empetr

ales Ephedrales Ericales EriocaulalesErythropalaless Escalloniales Eucommiales Euph

orbiales Eupomatiales Eupteleales Euryalaless

Fabales Fagales Falcatifoliales Ficales Flacourtiales Flagellariales Fouquieriales Franc

oales Frangulales

Galiales Garryales Geissolomatales Gentianales Geraniales Gesneriales Ginkgoales G

laucidiales Globulariales GnetalesGoodeniales Greyiales Griseliniales Grossulariales

Grubbiales Gunnerales Gyrocarpales Gyrostemonales

Haemodorales Haloragales Hamamelidales Hanguanales Heisteriales Helleborales He

lwingiales Himantandrales HippuridalesHomaliales Hortensiales Huales Huerteales

Hydatellales Hydnorales Hydrangeales Hydrastidales HydrocharitalesHydropeltidales

Hydrostachyales Hypericales Hypoxidales

Icacinales Illiciales Iridales Irvingales Iteales Ixiales

Jasminales Juglandales Julianiales Juncaginales Juncales

Lacistematales Lactoridales Lamiales Lardizabalales Laurales Lecythidales Ledocarpal

es Leitneriales LentibularialesLigustrales Liliales Limnanthales Linales Liriales Loasal

es Lobeliales Loganiales Lonicerales Loranthales LowialesLythrales

Magnoliales Malpighiales Malvales Marathrales Marcgraviales Mayacales Medusagy

nales Medusandrales MelanthialesMelastomatales Meliales Melianthales Meliosmal

es Menispermales Menyanthales Metteniusales MitrastemonalesMiyoshiales Moni

miales Moringales Musales Myricales Myristicales Myrothamnales Myrsinales Myrta

les

Najadales Nandinales Narcissales Nartheciales Nelumbonales Nepenthales Neuradal

es Nitrariales Nolanales NothofagalesNyctaginales Nymphaeales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

41

Ochnales Oenotherales Olacales Oleales Onagrales Oncothecales Opuntiales Orchid

ales Oxalidales

Saxifragales Pandales Pandanales Papaverales Paracryphiales Parasitaxales Paridales

Parnassiales PassifloralesPenaeales Pennantiales Pentaphragmatales Peridiscales P

etiveriales Petrosaviales Phellinales Philydrales PhyllanthalesPhysenales Phytolaccal

es Picramniales Pinales Pinguiculales Piperales Pittosporales Plantaginales Platanale

sPlumbaginales Poales Podocarpales Podophyllales Podostemales Polemoniales Pol

ygalales Polygonales PontederialesPortulacales Posidoniales Potamogetonales Primu

lales Proteales

Quercales Quillajales Quintiniales

Rafflesiales Ranunculales Rapateales Resedales Restionales Rhabdodendrales Rhamn

ales Rhinanthales RhizophoralesRhodorales Rhoipteleales Roridulales Rosales Rouss

eales Rubiales Ruppiales Rutales

Sabiales Salicales Salvadorales Samolales Samydales Sanguisorbales Santalales Sapin

dales Sapotales SarracenialesSaxegotheales Saxifragales Scheuchzeriales Sciadopitya

les Scleranthales Scrophulariales Scyphostegiales SedalesSilenales Simmondsiales S

milacales Solanales Sphenocleales Sphenostemonales Stangeriales Stellariales Stemo

nalesStilbales Stylidiales Styracales Surianales

Taccales Tamales Tamaricales Taxales Taxodiales Tecophilaeales Terebinthales Tern

stroemiales Theales TheligonalesThymelaeales Tiliales Tofieldiales Torricelliales Tov

ariales Tribelales Trilliales Trimeniales Triuridales TrochodendralesTropaeolales Turn

erales Typhales

Ulmales Urticales

Vacciniales Vahliales Vallisneriales Velloziales Veratrales Verbenales Viburnales Vinc

ales Violales Viscales VitalesVochysiales

Welwitschiales Winterales

Xanthorrhoeales Ximeniales Xyridales

Zamiales Zingiberales Zosterales Zygophyllales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

42

ALPHABETICAL LISTING OF ALL FAMILY NAMES OF SEED PLANTS WITH LINKS

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cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

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llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

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ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

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imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

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Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 11: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

11

relationships and applying it across all flowering plants would both be difficult and cause

substantial changes - see also below) Humphreys and Linder (2009) provide a well-

documented survey of generic concepts in plants which the reader should consult they

note that generic limits (broad versus narrow) have oscillated historically and that

currently larger genera tend to be recognised because studies tend to be on a broader

scale than in the past

Note that invoking similarity or difference - whether qualified (considerable

similarities substantial differences) or not - in a cladistic context as justification for

combining or splitting taxa is not a particularly strong argument (see eg

CardiopteridaceaeStemonuraceae - Karingrehed 2002c) Similarity and difference can

neither be defined precisely since what may seem to be substantial similarities to me

may not to the next person nor are they likely to be stable in the face of our changing

knowledge of morphology and what might be synapomorphies

I might have prefered to merge some families recognised here or split others but

by and large I do not think my own preferences matter very much - and I take the same

position with regards to comparable preferences expressed by others Indeed the

bottom line is that in flagged hierarchies of the kind used here the limits of any

monophyletic unit generally taught and discussed particularly other than species can

be established only by convention and consensus (eg Stevens 2002 2006a Entwisle amp

Weston 2005) This is rather different from the reasonable observation that the

phylogeny itself cannot be achieved by consensus (Thorne 1976) for in most cases there

is a fact of the matter when it comes to relationships Given the increasing support for

the outlines of angiosperm phylogeny a stable consensus classification based on this

phylogeny seems attainable Indeed in addition to providing current ideas of

relationships of seed plants in a synthesised form this site is part of an attempt to build

such a consensus about the circumscription of taxa (see APG 1999 2003 2009 Grass

Phylogeny Working Group 2001 Mabberley 2008) Reaching such a consensus is vital

since what we know of angiosperm phylogeny allows a very large number of

classifications to be based on it and as we find out more the number of possible

classifications increases greatly Unfortunately however nature does not dictate what

the classification should be All classifications are constructed by humans to

communicate particular aspects of groups and relationships Our goals as systematists

are surely to produce robust hypotheses of relationships to understand the evolution of

morphology and the like - but not to argue ad nauseam whether something should be

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

12

a family or a subfamily That way surely lies madness and worse the discredit of our

discipline

There are similar issues whatever naming system is used Thus in phylogenetic

naming (Baum et al 1998 for an example but cf Baum et al 2004 for the PhyloCode

see Cantino amp De Queiroz 2006) an unflagged hierarchy is used in which any

terminations of names used are uninformative about the relative position of taxa If one

adopts the principle of phylogenetic naming one indeed does not have to worry about

the nomenclatural consequences caused by lumping or splitting any well-supported

clade can be named without affecting the name of more or less inclusive clades

Unfortunately unflagged hierarchies have very serious deficiences as communication

devices because they lack one aspect essential in language biological or otherwise -

they contain no intrinsic information about the relationships of the group in question to

others (eg Pfeil amp Crisp 2005 Stevens 2006a) Recent suggestions for using prefixes like

Apo- and Pan- to PhyloCode names will however allow limited information of this

kind to be conveyed but only as it pertains to individual branches and current

proposals do not even mandate that the prefixes be employed consistently In any

event such proposals simply prevent the potential tripling of the number of quite

different names used to describe different aspects of a phylogenetic tree over those

used to name monophyletic groups pure and simple In general where n is the number

of extant species in a group the number of clades in such a group = n-1 (Species will

also need names too for their names see Dayrat et al 2008) Importantly here too

consensus over the clade names commonly learned by students and used in herbaria is

needed otherwise communication will be impeded the names themselves will provide

no guidelines as to which should be chosen The situation is of course more complicated

than this Terminations that convey ideas of rank in a phylogenetic classification can also

be used in phylocode names - however there they will carry no implications of rank

How they will be used is another matter of course

Of course there are other philosophies of classification and some still prefer

evolutionary classifications There classificatory principles differ substantially from those

followed here eg the recognition of paraphyletic taxa may be permitted however

detailed reasons for prefering the taxa that are recognised are rarely given although

nature and natural groups are often mentioned (cf Stuessy amp Koumlnig 2008) For

summaries of commonly used systems see Brummitt (1992) and Mabberley (2008)

new evolutionary systems appear every year or so However even those who allow or

promote the recognition of paraphyletic groups (eg Grant 2003 Thorne 2007

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

13

Heywood et al 2007) may find it of interest to examine a system recognizing only

monophyletic groups it provides a rather different understanding of evolution

Evolutionary classifications in general try and combine phylogeny and

morphological gaps although that is no easy thing to do - it is akin to combining chalk

and cheese (for an attempt to make this impossible task seem to be more objective see

Stuessy amp Koumlnig 2008) However there is a principle from evolutionary classification

that is relevant and I think quite useful here The size of the gap between two groups

tends to be inversely proportional to the sizes of the groups involved (Davis amp Heywood

1963) One can imagine a situation in which a large group is formally divided even

although the distinguishing characters of the two are weak whereas a smaller group

similarly divisable will be left intact

To summarize If hypotheses of phylogeny remain stable we should be able to base

a stable classification on that phylogeny and then get on with our work that is testing

the phylogenies we have elucidating phylogenies in areas where relationships are

unclear studying the evolution of morphology describing species etc In this context

the spread of the Angiosperm Phylogeny Group system (see below) and its widespread

utilisation in technical literature also floras (eg van der Meijden 2005) textbooks (eg

Simpson 2006 Judd et al 2007 [third edition]) dictionaries (Mabberley 2008) more

popular literature (eg Souza amp Lorenzi 2005 Spears 2006) and as an outline for a new

herbarium sequence (Haston et al 2007) is gratifying The posibility that one might be

able to develop a stable phylogeny-based classification of families and in particular

orders represents a dramatic turn-around from the pessimistic attitude about such

higher-level groupings expressed by Davis and Heywood (1963) and Thorne (1976) the

latter even suggesting that we should bury forever the metaphor of the phylogenetic

tree as highly unrealistic (ibid p 56) Returning to Godfray and Knapps (2004) users

of classifications who want a stable informative and accessible classification that

enables easy identification - unfortunately they want cake with everything and cannot

get it - these pages attempt to satisfy as many of their needs as possible but phylogeny

and monophyly are the primary shapers of the APG classification

On this classification in particular

Here I very largely follow the Angiosperm Phylogeny Group classification (APG

2003) Any differences are not to be interpreted as differences in principle simply that

new phylogenies continue to be published and here I attempt to provide an overview

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

14

of current ideas of higher-level relationships of seed plants The Angiosperm Phylogeny

Group classification is based on well-supported relationships evident in the numerous

molecular studies that began to appear in the late 1980s (see APG 1999 for the

principles underlying the classification) but recent phylogenetic work does not

contradict the major outlines of the trees used by APG II (2003) or even those in APG

I (APG 1999) The APG classification has for the most part been conservative and I

am similarly conservative here For the names and authorities I follow APG III (2009)

although the names of the authors may not always be correct these being in something

of a state of flux for the superordinal names used see Chase and Reveal (2009)

although classification at this level here is not exhaustive - only clades with several

orders have superordinal names

Some changes in our ideas of relationships and hence in the clades we talk about

are particularly likely in parts of Lamiales Caryophyllales and Malpighiales for example

although this will not affect the orders themselves Thus some changes to clade

circumscriptions are to be expected although I do not expect them to be substantial

but changes are neither a defect of cladistics nor a necessary consequence of the use of

molecular data Clades are hypotheses of relationships and as hypotheses they may be

overturned However work since APG I has not suggested other than minor changes

in the compositions of the orders even if the odd genus or even family is turning out to

be seriously misplaced - recent examples are Hydatellaceae (from monocots-Poales to

Nymphaeales Saarela et al 2007) Guamatelaceae (from Rosales-Rosaceae to

Crossosomatales Oh amp Potter 2006) and Perrottetia and Bhesa (from Celastrales-

Celastraceae to Huerteales and Malpighiales respectively Zhang amp Simmons 2006) The

main changes have been clarification of the relationhips of individual families or groups

of families that were of uncertain position eg of Chloranthaceae (Moore et al 2007)

Ceratophyllaceae (Jansen et al 2007) and unplaced asterid II families (Winkworth et al

2008a) As mentioned above phylogenies in the area of main interest in these pages are

overwhelmingly tree-like Although there is evidence for acquisition of host plant genes

by parasites (see below) even wider but inexplicable transfer of mitochochondrial

genes in plants like Amborella (Bergthorsson et al 2004 cf Goremykin et al 2009a)

and even transfers of nuclear genes (Vallenback et al 2008) these are the exception

rather than the rule

In cases where the Angiosperm Phylogeny Group II allowed alternatives as to the

limits of families - Papaveraceae in the broad sense or Papaveraceae plus

Pteridophyllaceae plus Fumariaceae Proteaceae in the broad sense or Proteaceae plus

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

15

Platanaceae - the choices made here follow common usage eg as in textbooks like

Judd et al (2007) and Simpson (2006) and particularly in the new edition of

Mabberleys The Plant Book (Mabberley 2008) This latter is itself an attempt to reflect a

consensus the result of taking the opinions of botanists at several meetings A largely

similar consensus is reflected in the APG III classification (APG 2009) where you will

see that alternative classifications have been dispensed with For many people the

existence of such alternative classifications will simply confuse so agreement over

which groupings to use when alternatives are permitted and then dispensing with the

alternatives that are not commonly used seems reasonable I am making the few

changes that are needed to bring the classification here in line with these others but

only gradually since there is not enough time in each day

As already mentioned higher-level relationships in general and the composition of

orders in particular have always presented something of a challenge to systematists

(eg Davis amp Heywood 1963) That being said the composition of some clades like

Apiales Crossosomatales and Pandanales is decidedly unexpected However it is

interesting to see that these higher level clades are generally accepted even in works

with different classificatory philosophies (eg Heywood et al 2007) For clades like

Malpighiales many of the family groupings within Asparagales etc attempts to find

distinctive characters have largely failed (but see Endress amp Matthews 2006a also

the Apomorphies page here) Interestingly as with families some groupings suggested

by molecular studies are supported by morphological andor chemical characters that

have long been known sometimes for over a hundred years the relationship between

Pittosporaceae and ApiaceaeAraliaceae are a case in point (Hegnauer 1969b and

references) As our knowledge of morphology and chemistry improves we can hope for

improvements in the characterisations of clades at all levels

In many cases the new family limits of the Angiosperm Phylogeny Group (see

APG 1999 2003) are not really controversial although changes from the limits

commonly accepted only a decade ago are sometimes quite dramatic (eg Wagenitz

1997) Thus the split of the old Saxifragaceae sl is necessitated by its extreme

polyphyly as also with Icacinaceaesl and Cornaceae sl However the limits and

relationships of such groups had long been considered to be unsatisfactory but until

recently there had been no real reason to prefer one proposed arrangement over

another It is generally accepted that the limits of Lamiaceae and Verbenaceae have to

be redrawn although there is as yet no compelling evidence that the redrawn taxa are

not sister taxa (there is no evidence that they are) But whatever their relationships the

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

16

content of the clades has changed considerably and incidentally they are now easier to

identify than before the decision to recognise two families is not difficult The same is

true for Salicaceae and Achariaceae (Malpighiales) two previously small families that

have received the bulk of the old Flacourtiaceae Clade and hence taxon limits remain

difficult around Theaceae (Ericalessee alsoSladeniaceae Pentaphylacaceae)

and Euphorbiaceae (Malpighiales see

also Phyllanthaceae Picrodendraceae Putranjivaceae etc) Here current groupings

may not represent quite such dramatic changes in our understanding of relationships

Indeed parts of the old Euphorbiaceae that were separated may yet go back together

particularly Phyllanthaceae and Picrodendraceae (see Wurdack et al 2004 Davis et al

2005) even if Theaceae should indeed be dismembered (Geuten et al 2004) However

given that there is no molecular evidence that currently warrants combining all the

segregates of Euphorbiaceae and even if some do come together the clades the

families represent suggest novel groupings not recognised in current classifications

Furthermore Rafflesiaceae appear to be embedded within Euphorbiaceae s str (Davis

et al 2007) so the family is divided Of course maintaining even a moderately narrowly-

circumscribed Euphorbiaceae would entail reducing the iconic Rafflesiaceae to

synonymy Relationships in core Caryophyllales especially

around Phytolaccaceae andNyctaginaceae and also Portulacaceae are incompletely

understood and refashioning of taxon limits will doubtless be needed as cladistic

relationships become apparent (eg see Nyffeler 2007 Ogburn 2007a b) Some

groupings in the old Icacinaceaeand Olacaceae also remain unclear

The discovery of the relationships of parasitic and aquatic groups have presented a

particular challenge to systematists Morphologically some of these plants are so highly

modified that interpretation of the plant body in conventional terms is difficult or even

impossible Thus parasitic groups (broad-leaved angiosperms only) such as Rafflesiaceae

are hard to place since both the vegetative body and the flowers are changed almost

beyond recognition (flowers of rafflesiaceae are a good example of gigantism - Davis et

al 2007) Furthermore plastid gene sequences may be difficult or impossible to obtain

the chloroplast DNA in particular being highly degraded and the rate of molecular

change in general may be high so the problem of long-branch attraction is serious (eg

Duff amp Nickrent 1997 Nickrent et al 1998 Caddick et al 2002a G Petersen et al

2006b) Echlorophyllous sapromycoheterotrophic taxa mostly known from monocots

present similar problems However progress is being made placements for

Apodanthaceae Rafflesiaceae Mitrastemonaceae Cytinaceae and Cynomoriaceae have

recently been suggested (Barkman et al 2004 2007 Davis amp Wurdack 2004 Nickrent et

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

17

al 2004 Davis et al 2007) it seems likely that Burmanniaceae sl are polyphyletic but

both parts are in Dioscoreales (Merckx et al 2006) and relationships within the largely

hemiparasitic Santalales are also gradually being clarified (Maleacutecot 2002) The inclusion

of parasitic taxa in general molecular analyses can cause conniptions (eg Nickrent et al

2004 Davis et al 2004 Chase et al 2006 G Petersen et al 2006b) and there can be

horizontal transmission of genes (eg Davis amp Wurdack 2005 Vitaceae to Rafflesiaceae

Barkman et al 2007 the mitochondrial atp1 gene commonly moves)

Similarly in water plants neither vessels in particular nor much xylem in general is

needed leaves are highly modified and water-mediated pollination if adopted may

well be associated with major changes in floral morphology Here too recent molecular

studies suggest that aquatic groups with hitherto problematic relationships may find

homes Thus Podostemaceae are close to Clusiaceae and Calophyllaceae and are sister

to Hypericaceae (Malpighiales Kita amp Kato 2001) Hydatellaceae which used to be in

Poales are part of Nymphaeales (Saarela et al 2007) and Hydrostachyaceae may be

close to Hydrangeaceae (Cornales Xiang et al 2002) or perhaps should be placed in

Lamiales note that in the first two cases in particular there are morphological and

chemical features that support such a move If Podostemaceae are indeed close to

Clusiaceae I look forward to seeing hypotheses to explain how the dramatic changes in

the vegetative body that have made Podostemaceae so problematic for generations of

systematists took place That conventional wisdom has trouble in understanding or

explaining how the morphologies of groups like Clusiaceae and Podostemaceae can be

related is largely a problem with conventional wisdom Ceratophyllaceae seem to be

finding a position as sister to eudicots (see Moore et al 2007) but their morphology is

so derived that there is no morphological evidence of which I am aware for this

relationship There are also similar although less extreme morphological problems with

many plants pollinated by wind

In such situations especially with some water plants mycoheterotrophs and

parasites a variety of characters may be affected and particularly under such

circumstances groups have tended to be formed using a few or even only a single

character that seems to provide evidence of relationships Add to this the tendency to

weight some characters particularly strongly on a priori grounds and the result was

either the recognition of conglomerate taxa such as Amentiferae which now appear to

be highly polyphyletic or the segregation of families like Plantaginaceae s str (now

much expanded) and Leitneriaceae (now in Simaroubaceae) - all of these are more or

less wind pollinated

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

18

Even although relationships of many parasitic and aquatic groups are being

clarified and at least some of the proposed phylogenetic positions seem to be well

supported it is indeed often still very difficult to understand how such groups have

evolved from their more morphologically conventional relatives However we have had

all sorts of preconceptions as to how characters should evolve Thus we used to

assume that features like highly scalariform vessel perforation plates or the complete

absence of vessels or a flower with an androecium that had many stamens a superior

ovary or separate petals were necessarily plesiomorphic or primitive and conversely

a simple perforation plate and the presence of vessels an androecium with few

stamens inferior ovary or petals that were connnate were almost necessarily

apomorphic or advanced (but cf eg Stebbins 1951) Such assumptions are incorrect

(eg Soltis et al 2005b) Carpels may become secondarily free carpels may fail to close

the seeds then developing outside the confines of the carpel as in some Aspagaraceae-

Nolinoideae Violaceae Berberidaceae Malvaceae-Sterculioideae etc) in Peliosanthes

teta perhaps the only species in Peliosanthes(Asparagaceae-Nolinoideae) the ovary

varies from superior to inferior (Jessop 1976 species limits here need close

investigation see also Kuzoff et al 2001 and Soltis amp Hufford 2002 Saxifragaceae

Apiales etc) many-seeded carpels can evolve from few-seeded carpels

(Razafimandimbison et al 2008) monoecy may be derived from dioecy (Schaefer amp

Renner 2010 and references) Classic studies such as those by Babcock (eg 1947)

on Crepis that assumed that evolution - in this case of the karyotype in particular - was

unidirectional have needed comprehensive re-evaluation (Enke amp Gemeinholzer 2008)

Most if not all characters have reversed andor evolved in parallel as is clear in the

discussion of ovary position here especially in Poales Saxifragales and Asparagales and

also in comments on such morphologically distinctive taxa (when compared with their

immediate relatives) such as Menyanthaceae and especially Pittosporaceae Parallel

evolution may occur even at the level of amino acid substitution as in the independent

acquisition of the phosphoenolpyruvate carboxylase (pepC) gene in C4 photosynthesis in

grasses (Christin et al 2007b see also Blaumlsing et al 2000)

Thinking of how such characters have been used in the past in classifications at the

generic level the results have been similar to those just discussed above for parasitic

plants and others Relying too much on animal pollination syndromes to mark generic

boundaries has all too often led to taxa that are highly unsatisfactory phylogenetically

(see eg Acanthaceae Bignoniaceae Campanulaceae Ericaceae Lamiaceae

Orchidaceae) and over-reliance on characters of fruit and seed (see eg Brassicaceae

and Apiaceae) has also led to unsatisfactory generic limits Again the more general

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

19

problem is the use of one or a very few characters that have been weighted a priori to

structure classifications (see also Garciacutea et al 2009) although it is perhaps not

surprising that such an approach sometimes results in the delineation of taxa that have

indeed turned out to be quite distinct (cf eg Burtt Astragalus versus Oxytropis)

Taxa recognized here are monophyletic yet many are polythetic at the

morphological level that is they lack unique features characterizing (defining) all and

only members of that family they can be recognised phenetically only by the unique

combinations of characters that they posess This is the result of evolution any

synapomorphy characterizing a taxon may be lost or modified beyond easy recognition

in some of its members or the synapomorphy may appear to be identical to a feature

that has evolved in parallel in a quite unrelated plant That plant groups are polythetic is

almost as much a feature of monophyletic taxa as the paraphyletic taxa common in

evolutionary classifications Some families now include substantial variation as

phenetically distinct derived groups are placed in their proper phylogenetic position -

examples are Ericaceae which include the erstwhile Empetraceae a wind-pollinated

group that is florally very different and the overwhelmingly large-flowered and animal-

pollinated Plantaginaceae which now include several derived small-flowered aquatic

and wind-pollinated groups that were previously placed in separate families

Thus some families as delimited here may not be easy to recognize However

remember that detecting relationships - use whatever characters you can even if they

are not obvious - and naming a plant - focus on easy-to-see characters that may not

reflect relationships - are quite different problems Taxa although natural may not be

readily recognizable indeed it was in exactly this context that Lamarck worked out the

basic principles of writing dichotomous keys in 1778 (see Scharf 2007 for keys and the

like) Of course Lamarcks idea of nature was very different from ours - he thought that

there was some kind of continuum of form on which living organisms were to be

situated with no real gaps anywhere - but this meant that his genera (for example)

might well not be sharply distinct from each other even if each were part of the real

continuum that was life Lamarcks separation of the establishment of relationships and

groupings in a system and identification of the plants in that system have been widely

accepted (but cf eg Godfray amp Knapp 2004 many of the contributors to Heywood et

al 2007) Perhaps the best way of identifying plants at the family level is by well-made

multiple access keys as in Watson and Dallwitz (1992a onwards family limits there may

differ substantially from those adopted here) Multiple access keys free users from the

constraints of dichotomous keys in which particular characters are needed at each step

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

20

of the identification process before there can be further progress Instead those

characters that happen to be evident on a specimen can be used in whatever order is

convenient when linked to illustrations glossaries etc their power is enormous (see

Dallwitz et al 2000 [2006] for the principles underlying their construction and use)

Nevertheless dichotomous keys such as those of Hutchinson (1973) and Franz Thonner

(Geesink et al 1981) have their uses Of course taxa coming out adjacent in keys may

well not be at all related

(When identifying large numbers of plants even more efficient than either style of

identification and certainly lots more fun is sight identification Unless you have a

photographic memory you have to build up your knowledge of comparative plant

morphology - on which the ability to make accurate identifications depends - by

repeated observation When faced with an unknown plant I always look for leaf teeth

and stipules distinctive hair types especially stellate and T-shaped hairs glands and

punctations of any sort the presence of latex or other exudate and check leaf insertion

smelling crushed leaves can also be helpful In this context nodal anatomy can usually

be checked using a razor and a hand lens or even simply carefully examining leaf scars

The short paragraphs added after most families may help in confirming familial

identifications)

For the record and for the little that it is worth there are 4 orders and 13 families

of gymnosperms characterised on these pages and together they include some 82

genera and 947 species For angiosperms comparable figures are 56 orders 445

families 13208 genera and 261750 species (of which monocots include 11 orders 89

families 2759 genera and 52760 species) Note however that higher mathematics

was never my strong point and anyway these are pretty meaningless figures even for

species which many (but not all) might concede smacked slightly more of reality than

other taxa estimates range as high as 422000 (Govaerts 2001) Furthermore numbers

of genera and species change daily Nevertheless as emphasized here families are

useful in teaching we as a community can ensure that their limits remain largely stable

and by concentrating on relatively few of them one can gain some familiarity with much

of the worlds flora For summaries of the sizes of orders and families and of the general

arrangement followed in these pages see also the Statistics page itself

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

21

A PHYLOGENETIC TREE OF SEED PLANTS (THE MAIN TREE)

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

22

The Main Tree above is a conservative summary of well supported relationships

between the major clades of extant seed plants Reading - interpreting and

understanding - such a phylogenetic tree is not necessarily easy so reading a tree may

require a bit of practice OHara (eg 1988 1992) provides an accessible introduction to

thinking and talking about history in the context of looking at phylogenetic trees More

recently Krell and Cranston (2004) Crisp and Cook (2005) and others have emphasized

how careful one must be when interpreting and talking about evolution in ladderized

trees in particular and phylogenetic trees in general Gregory (2008) provides a

particularly detailed discussion of this problem In ladderized trees the smaller (in terms

of numbers of terminals) sister taxon is consistently shown on the same side at every

node thus the trees tend to be pectinate like the teeth of a comb however the

horizontal axis that is as a result so evident has no polarity or particular direction The

use of the adjective basal when discussing phylogenetic trees is especially dangerous

(see also Wojciechowski et al 2004 D Soltis et al 2005b) When I use the term and the

context is not otherwise clear I am referring to the pectinations at the base of a

ladderised tree Remember that when talking about sister taxa one can never be basal

to the other or older or younger although members of one can be more derived - in the

sense of having more apomorphies - than the other (but even then one has to be

careful) The words primitive and advanced and lower and higher should

practically never be used especially when talking about taxa Amborellaceae and

Pinaceae are sister to all other flowering plants and Pinales and are basal to the crown

groups of all other flowering plantsPinales respectively but that does not mean they

are thereby primitive Pinaceae in particular have numerous apomorphies The word

plesiomorphic is far less loaded than primitive and can be used to talk about

individual characters

Most trees here have been more or less ladderized as is the one above The

ladderization may be imperfect for example you can see that on the Main Tree

asterids with ten orders follow rosids and relatives which have seventeen although

this is in part because of the currently unresolved nature of relationships between the

main clades of core eudicots (this is likely to change soon) The Main Tree could have

been drawn with Amborella Acorus or a host of other taxa at the far right without

offending any relationships phylogenetic trees are like mobiles the only fixed points

being the nodes Despite the problems just mentioned pectination interpreted

carefully has its value and it is not an arbitrary process As one reads the terminals of a

pectinate tree from left to right adjacent terminals will be separated by apomorphies

that will cumulate in an appropriate fashion When thinking of a book or a herbarium

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

23

sequence (see Haston et al 2007) this is of some value Nymphaeales and

Austrobaileyales are here adjacent on the tree but they could be separated by

hundreds of families in the sequence merely by rotating only the node from which

Nymphaeales and their sister group (all other angiosperms minus about 75 species)

arise If adjacent in a book or herbarium or as in these summary pages then it is

relatively easy to relate their characters both apomorphic and plesiomorphic but if

separated by hundreds of pages or two floors in a large building then it is less easy to

get anything from the sequence (Since all orders in the pages of the main part of this

site are preceded by the apomorphies of all nodes immediately below them in the seed

plant phylogeny and because of the linkages that have been built in to the site this

problem is much less serious there there is no sequence) So the particular sequence of

all seed plants that is based on a phylogenetic classification but is to be used in herbaria

or any other place where taxa have to be arranged linearly as in this part of APweb

will be one that maximizes the number of taxa that are both successive branches of the

tree and placed successively in the sequence Since specimens are generally filed under

families the outline of a new family sequence for arranging herbaria and books can now

be suggested (Haston et al 2007) and such a standardized sequence will further help

teaching and learning about plants

ON THINKING ABOUT APOMORPHIES

Identifying apomorphies is important because understanding the evolution of

morphology in the broad sense - ie including anatomy chemistry etc - in the context

of diversification is one of our major goals For identifying apomorphies several

preconditions must be met One needs to have an accurate well supported phylogeny

one has to have examined the right taxa both from the point of view of morphology and

molecules one has to have coded the characters right (ie delimited states

appropriately) and one has to use an appropriate model of evolution when fitting the

variation to the tree (see eg Omland 1999 Stevens 2006b) If the distributions of

apomorphies below are compared with those in other studies (eg Turgeon et al 2001

Bremer et al 2001 Endress 2001 Albach et al 2001a Judd amp Olmstead 2004 D Soltis

et al 2005b Zhang et al 2006 Judd et al 2007) the reader may find a number of

differences Although I have integrated such studies as far as is possible there are five

reasons that link to the preconditions just mentioned why there may be differences

where features are placed on trees

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

24

1 I may not have found all the information about a particular character there may be

disagreement over its interpretation or I have added information

2 The sampling of nearly all molecular studies is very incomplete (see Salisbury amp Kim

2001 for problems caused by sampling) indeed it is commonplace to decry the

incompleteness of molecular sampling But not only is the sampling in molecular studies

often less than we might wish that of the morphological and chemical characters whose

evolution we are interested in understanding is also often very poor So for many

anatomical chemical and embryological characters that are confidently said to

characterise families and other groups we all too often have no idea if those characters

are applicable to the whole clade or just to a subgroup within that clade However as

our sampling improves we can locate changes on the tree more precisely if in

unexpected positions Thus is is sometimes suggested that Ericaceae have ellagic acid

(Soltis et al 2005b) in fact the little evidence we have indicates that only a small clade

of some 80 species has ellagic acid and the rest of the family - itself alone about one

third of all Ericales - does not Similarly Ericaceae are noted for their

ectendomycorrhizal associations but Enkianthus sister to all other Ericaceae

apparently lacks these (Abe 2005) Thus these mycorrhizae and their associated

distinctive hair roots are unlikely to be apomorphies of Ericaceae - however it appears

that other families near Ericaceae may also have ectendomycorrhizae (see Asai 1934)

3 I am fitting characters to a very conservative tree with many polytomies although the

nodes that are utilised are for the most part strongly supported polytomies make the

optimisation of characters that is the assigment of character state change to a

particular node on the tree notably difficult (eg Madison amp Madison 2002) In nearly

all studies of the evolution of characters distributions of characters are optimised on a

more or less fully resolved tree and the construction of supertrees may yield yet more

detailed hypotheses of relationships (for literature on supertrees see Cotton amp

Wilkinson 2007 2008) Of course some nodes on such fully resolved trees andor

supertrees may have little support and optimisations of characters on such trees may

carry correspondingly little conviction Even parts of some of the trees used here have

poor support eg relationships within aquatic Alismatales etc although I indicate

when this is the case

4 Exactly how one goes about optimising a character on a tree is critically important

Even using simple parsimony optimisations (ACCTRAN or DELTRAN ACCelerated

TRANsitions or DELayed TRANsitions) the position of synapomorphies on trees - and

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

25

hence our ideas of evolution - may differ and this is still more true if one uses maximum

likelihood or Bayesian methods and makes apparently reasonable suggestions about

weighting gains over losses (eg Donoghue amp Ackerley 1996 Cunningham et al 1998

Omland 1997 1999 Ree amp Donoghue 1999 Polly 2001 Webster amp Purvis 2001

Ronquist 2004 Crisp amp Cook 2005) Sannier et al (2007) show how in in Arecaceae that

where on a phylogenetic tree one might peg changes in microsporogenesis will depend

on the methods one uses to do this and Pedersen et al (2007) discuss the sometimes

very substantial effect of node support on the posterior probabilities of ancestral

character states Here I use parsimony optimization not always as explicit as it might be

but I have often indicated where there are particularly important uncertainties as to the

positions of particular character changes on the tree

5 Finally although I have paid quite a lot of attention to the delimitation of the

character states that make up all the characterizations I have not spent enough time on

this critical operation If we are interested in understanding evolution then fitting the

basic variation - not character states - to a tree in principle allows greater flexibility in

understanding morphology in the context of local phylogenies (see also Stevens 2000

Endress 2005c) However many character states used here are delimited globally that

is they are circumscribed in the context of the variation shown by individual characters

across all angiosperms andor in the context of classic ideas of character evolution

Character states often have arbitrary limits and serve best to communicate

information whether they are in fact suitable for either phylogenetic analysis or

understanding evolution are separate issues Studies have rather unsurprisingly

perhaps but importantly shown that dividing the one character into different sets of

states may yield differing ideas of evolution of that character (eg Lamb Frye amp Kron

2003 Hibbett 2004) When looking at trees on which character states are optimised

one should bear in mind the problems surrounding the delimitation of states (eg

Stevens 2000 2006b) and the danger of using pollen or other types - constructs

based on many characters that vary independently but which effectively get lost in

these types thus Blackmore et al (2009) decomposed the pollen types in Asteraceae

into 52 characters

I use Remanes three main criteria of homology or better similarity when

determining the basic similarity of structures on different organisms (see Remane 1952)

These criteria are special properties position and intermediates Special properties

include anatomical or chemical characters gene expression data etc Position refers

to the position of an organ with respect to landmarks on the plant Although plants are

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

26

plants and landmarks are not as fixed as one might like trying to work out the relative

positions of parts is a good way of understanding morphology so long as one does not

try to out-do Procrustes Intermediates include intermediates found on the same

plant or on different plants Intermediates may observed during development or be

adult structures Thus one may find changes during development which suggest what

the nature of a particular structure is and very different stuctures on different plants

may turn out to be similar early in development Or one can simply compare different

taxa and see that structures that appear to be very different when comparing two

different taxa can be linked morphologically by looking at other taxa

Nevertheless the use of these criteria may not yield an unambiguous answer as to

what a structure is even given a solid phylogeny and an improved understanding of

development (see Jaramillo amp Kramer 2007 for a useful discussion) As Endress (2005c)

observed a number of features - position function development shape anatomy

histology gene activity and relationships to other taxa that clearly have petals - can be

used to distinguish a petal (for example) from other floral structures if a petal does not

have one of these features is it thereby not a petal Thus Maturen et al (2005) recently

found that floral organ diversity genes (B and C) were expressed in the large white

inflorescence bracts of Cornus (see also Costa et al 2005) Peney et al (2005) noted that

not all monosulcate pollen grains in monocots have the same developmental pathway

and that as a result such pollen might not have the same ancestral state Reeves and

Olmstead (2003) suggested that the genetic mechanisms causing monosymmetry in

Lamiales and Solanales were different and Serna and Martin (2006) described similar

problems with the development of hairs in Arabidopsis when compared with that of

hairs in Antirrhinum and Solanaceae Indeed as one perhaps might expect delimitation

of states and characters does not necessarily become easier with increasing knowledge

of development etc Thus Buzgo et al (2004) Matthews and Endress (2005) and others

have shown how hard it can be to distinguish between eg prophylls and other floral

structures as their behaviour is studied during the course of the development of the

flower Similarly at what concentration is a particular secondary metabolite deemed to

be present (Waterman (2007)

The validity of the approach used here that of fitting morphological variation to a

largely molecular-based tree may be questioned However I think it rather unlikely that

well-supported molecular branches will be overturned by morphological data Indeed

analyses of morphological data alone do provide support for many of the clades evident

in molecule-only analyses and analyses with morphological and molecular data

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

27

together may lead to increased support for clades (eg Hufford 1992 Nandi et al 1998

[but adding morphological data reduces support for a number of critical clades too]

Doyle amp Endress 2000 however in none of these papers is the use of morphology

without ambiguity) It is unfortunately clear that the use of morphology alone may not

suggest problems in the phylogenetic placement of taxa that later turn out to have been

wrongly included (eg Zhang et al 1992) Thus I have been wary of putting much weight

on clades that have only morphological support but note that I have treated molecule-

based clades with low bootstrap or jacknife support values (esp below 70) or low

posterior probabilies (below 095) likewise Although I may have been mistaken in

placing so much emphasis on molecular data in terms of providing the basic

phylogenetic framework for angiosperms morphological and molecular data are only

very rarely in irreconcilably strong conflict There are indeed a few places where the

conflict seems extreme These include the relative positions of the Monimiaceae and

Hernandiaceae (Laurales) the position of Hanguanaceae (Commelinales [as here] or

Zingiberales) and of Triplostegia (is it in Dipsacaceae or Valerianaceae - see

Dipsacales) Fossils are unlikely to affect the topologies of the trees presented here but

see below for their importance in understanding morphological evolution in general and

the evolution of angiosperms in particular Nevertheless some largely reject the idea

that trees based on molecular data alone can recover phylogenetic relationships

especially when branching points are old and prefer to used trees based on analysis of

morphological data including those taken from fossils (Hilton amp Bateman 2006 Farjon

2007)

All in all however the extent of the congruence between morphological and

molecular data is impressive and heartening and many clades can be characterised

morphologically It seemed in 1998 that there were no unambiguous morphological

synapomorphies for angiosperm orders (K Bremer 2000) and this is still true if by

unambiguous is meant non-homoplasious However many orders and other clades

have synapomorphies even if these may be indistinguishable at least at the current

level of morphological and developmental knowledge from parallel occurences

elsewhere As our morphological knowledge increases so too does the number of

apomorphies

Endress and Matthews (2006a) emphasize the importance of tendencies and

developmental constraints when thinking about characters of clades within the rosids -

many characters or character combinations occur in a rather sporadic fashion within

the clade and apparently notably less frequently outside it There are a number of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

28

examples of tendencies here and in the individual order pages such as the features

enclosed by parentheses in groups above the level of family eg see (cuticular waxes

as aggregated rodlets) for the commelinids and the discussion of the distribution of

polyandry that is flowers with numerous stamens in the asterid I + II groups etc

Tendencies may involve more than single characters When thinking of the

associations of features like integument thickness and vascularization of the integument

(Endress amp Matthews 2006a) one can imagine fairly simple developmental

preconditions being operative As Endress and Matthews (2006a) note it is difficult to

imagine vascular tissue developing in an integument only two cells thick Changes here

would be loosely correlated if morphologically linked However the strongly correlated

changes noted by Givnish et al (2005) are ecologically linked but are presumably

morphologicallydevelopmentally independent When there seem to be characters

evolving more or less together tests can be carried out to see if the changes are

concentrated on certain branches of the tree (eg Maddison 1990 Sanderson 1991

Maddison amp Maddison 2000) Such tests have rarely been carried out at higher levels in

angiosperms

One of the most striking examples of a tendency is the distribution of N-fixation

restricted as it is to a monophyletic group of four clades although it has arisen seven (or

perhaps several more) times independently within the N-fixing clade and several

members of two quite different kinds of bacteria are involved (eg Clawson et al 2004

Elliott et al 2007 Sprent amp James 2007) Other examples of work that bears on the issue

of tendencies include the findings that flowers of polysymmetrical Arabidopsis have

genes like TCP1 that are expressed asymmetrically during early development and TCP1

is a probable orthologue of the well-known CYC gene of Antirrhinum that is involved in

the development of monosymmetric slowers there (Cubas et al 2001 Costa et al 2005

etc) Parallelism might build on this underlying morphologically cryptic monosymmetry

even if details of the genetic mechanisms causing the monosymmetry evident in

particular groups may be different (Reeves amp Olmstead 2003 Cubas 2004 see above)

Similarly there have been several recent suggestions that the capability to synthesise a

particular metabolite may be switched off but not lost and so can sometimes be

reacquired (eg Wink amp Witte 1983 Wink 2003 Liscombe et al 2005 Larsson 2007

Waterman 2007) Hence perhaps the rather spotty distribution of many secondary

metabolites like ellagic acid the indole alkaloid camptothecin iridoids etc when

considered in the context of phylogenies As a non-botanical example - but a rather nice

one - Salwini-Plawen and Mayr (1961) suggested some time ago that there has been

considerable parallelism (40-65 or more independent origins) in the evolution of eyes in

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

29

metazoans However Pax 6 seems to be a master control gene that is involved in eye

formation perhaps throughout all bilateralians (eg Gehring amp Ikeo 1999 Erwin amp

Davidson 2002) which changes the nature of the problem Note however that in

general our invocation of evolutionary tendencies to explain the patterns of

distributions of characters is really an admission that we do not understand those

patterns

As we find out more about variation we will probably find fewer and fewer features

constant throughout a group Most unqualified statements of presence and absence

should properly be qualified as usually present or usually absent if one is thinking of

the characterisations on the order pages as encompassing the total variation within a

clade Thus Pistia alone among monocots as so far known has sieve tube plastids with

starch grains not protein crystals However this does not affect the fact that sieve tube

plastids with cuneate protein crystals and lacking starch grains are an apomorphy for

monocots Furthermore fossils in a number of cases suggest character combinations

unknown in extant taxa as may be seen in the discussions of Fagaceae Platanaceae

Iteaceae Calycanthaceae etc Confusing the issue there may be questions as to where

exactly on the tree a particular fossil is to be placed (eg see Nymphaeaceae

Calycanthaceae Archaefructus etc)

To summarize given our current understandings of both phylogenies and

characters evolution of some characters in which we are interested seems very labile

(see eg D Soltis et al 2005b Endress and Matthews 2006a Stevens 2006b Ekman et

al 2008 for a good recent study) and I have been cautious when talking about character

evolution Much effort must continue to be spent in summarizing characters of well-

established clades at all levels providing features by which they may be recognized and

signaling synapomorphies Remember that (1) the basic morphological anatomical and

chemical knowledge of many critical taxa is woefully incomplete (2) different

assumptions about character evolution may greatly affect the position of

synapomorphies on trees (3) in many cases relationships within and between many

groups are too uncertain at present to worry very much about synapomorphies and

(4) we must be clear about what we do and do not not know As mentioned above I

have indicated in a number of places where there are particularly important

uncertainties as to where characters should be placed on the tree Nevertheless it is a

relatively easy matter to update notes such as these and it can be a simple matter to

incorporate new data on characters that have never before been considered in the

context of a tree Much basic - and unfortunately perhaps unfashionable - work must be

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

30

carried to clarify the distribution of morphological anatomical and chemical characters

for examples of what can to be done see the work of P K Endress and collaborators

(floral morphology and development) and S R Jensen and collaborators (iridoids) But

acquiring information about nodal anatomy cell and tissue distribution nodal anatomy

and the like is not difficult at all and should be generally encouraged

But all this is merely a necesary prelude to the understanding of evolution For this

one needs to know a lot more including dating the phylogeny understanding the

function(s) of characters factoring in relevant aspects of the palaeoenvironment in

which an apomorphy first appeared etc

SUMMARY OF THE SYSTEM

Below is a formalised summary of the relationships within orders of the families of seed

plants There are a few families that are not recognised even as options in APG II and

vice versa as well as a few extra orders The families to be recognised in the most

recent edition of Mabberleys The Plant Book (Mabberley 2008) are also largely

consistent with those below But all differences are trivial and will - I hope - eventually

disappear

Square brackets - [] - enclose clades the plus sign - + - designates sister taxa a

comma - - denotes part of a polytomy and quotation marks - - denotes a

paraphyletic group

SEED PLANTS

GYMNOSPERMS

Cycadales

Cycadaceae + Zamiaceae

Ginkgoales

Ginkgoaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

31

Pinales

Pinaceae [[Araucariaceae [Podocarpaceae + Phyllocladaceae]] [Sciadopityaceae

[Taxaceae + Cupressaceae]]]

Gnetales

Ephedraceae [Gnetaceae + Welwitschiaceae]

ANGIOSPERMSFLOWERING PLANTS

Amborellales

Amborellaceae

Nymphaeales

Hydatellaceae [Cabombaceae + Nymphaeaceae]

Austrobaileyales

Austrobaileyaceae [Schisandraceae + Trimeniaceae]

Chloranthales

Chloranthaceae

MAGNOLIIDS

Magnoliales

Myristicaceae [Magnoliaceae [[Himantandraceae + Degeneriaceae] [Eupomatiaceae +

Annonaceae]]]

Laurales

Calycanthaceae [[Siparunaceae [Gomortegaceae + Atherospermataceae]] [Monimiaceae

[Hernandiaceae + Lauraceae]]]

Canellales

Canellaceae + Winteraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

32

Piperales

[Hydnoraceae + Aristolochiaceae] [Piperaceae + Saururaceae]

MONOCOTS

Acorales

Acoraceae

Alismatales

Araceae [Tofieldiaceae [[Alismataceae [Hydrocharitaceae + Butomaceae]]

[Scheuchzeriaceae [Aponogetonaceae [Juncaginaceae [Maundiaceae [[Posidoniaceae

[Ruppiaceae + Cymodoceaceae]] [Zosteraceae + Potamogetonaceae]]]]]]]

Petrosaviales

Petrosaviaceae

Dioscoreales

Nartheciaceae [[Taccaceae + Thismiaceae] [Burmanniaceae + Dioscoreaceae]]

Pandanales

Velloziaceae Triuridaceae Stemonaceae [Pandanaceae + Cyclanthaceae]

Liliales

Corsiaceae [Campynemataceae [Petermanniaceae [Colchicaceae + Alstroemeriaceae]]

Melanthiaceae [[Philesiaceae + Rhipogonaceae] [Smilacaceae + Liliaceae]]]

Asparagales

Orchidaceae [[Boryaceae [Blandfordiaceae [Lanariaceae [Asteliaceae + Hypoxidaceae]]]]

[[Ixioliriaceae + Tecophilaeaceae] [Doryanthaceae [Iridaceae [Xeronemataceae

[Xanthorrhoeaceae [Alliaceae + Asparagaceae]]]]]]]

COMMELINIDS

Unplaced

Dasypogonaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

33

Arecales

Arecaceae

Poales

[Typhaceae + Bromeliaceae] [Rapateaceae [[Thurniaceae [Juncaceae + Cyperaceae]]

[[Mayacaceae [Eriocaulaceae + Xyridaceae]] [Flagellariaceae [Anarthriaceae

[Centrolepidaceae + Restionaceae]]] [Joinvilleaceae [Ecdeiocoleaceae + Poaceae]]]]]]]

Commelinales

[Commelinaceae + Hanguanaceae] [Philydraceae [Haemodoraceae + Pontederiaceae]]

Zingiberales

Musaceae [Strelitziaceae + Lowiaceae] Heliconiaceae [[Cannaceae + Marantaceae]

[Costaceae + Zingiberaceae]]

Ceratophyllales

Ceratophyllaceae

EUDICOTS

Ranunculales

Eupteleaceae [Papaveraceae [[[Lardizabalaceae + Circaeasteraceae] [Menispermaceae

[Berberidaceae + Ranunculaceae]]]]

Sabiales

Sabiaceae

Proteales

Nelumbonaceae [Platanaceae + Proteaceae]

Trochodendrales

Trochodendraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

34

Buxales

Haptanthaceae [Buxaceae + Didymelaceae]

CORE EUDICOTS

Gunnerales

Gunneraceae + Myrothamnaceae

Dilleniales

Dilleniaceae

Saxifragales

Peridiscaceae [[Paeoniaceae [Altingiaceae [Hamamelidaceae [Cercidiphyllaceae +

Daphniphyllaceae]]]] [[Crassulaceae [Aphanopetalaceae [Tetracarpaeaceae

[Penthoraceae + Haloragaceae]]]] [Iteaceae [Grossulariaceae + Saxifragaceae]]]]

Cynomoriaceae unplaced

Vitales

Vitaceae

ROSIDS

FABIDROSID I

Zygophyllales

Krameriaceae + Zygophyllaceae

Celastrales

Lepidobotryaceae + Celastraceae

Oxalidales

Huaceae [[Connaraceae + Oxalidaceae] [Cunoniaceae [Elaeocarpaceae [Brunelliaceae +

Cephalotaceae]]]]

Malpighiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

35

[Achariaceae [Goupiaceae [Violaceae + Passifloraceae] [Lacistemataceae Salicaceae]]

[Lophopyxidaceae Putranjivaceae] [Ctenolophonaceae [Erythroxylaceae +

Rhizophoraceae]] Linaceae Ixonanthaceae Humiriaceae Irvingiaceae

Centroplacaceae Pandaceae [[Bonnetiaceae + Clusiaceae] [Calophyllaceae

[Hypericaceae + Podostemaceae]]] [Malpighiaceae + Elatinaceae] Ochnaceae

[Peraceae [Rafflesiaceae + Euphorbiaceae]] [Phyllanthaceae + Picrodendraceae]

[Balanopaceae [[Trigoniaceae + Dichapetalaceae] [Chrysobalanaceae +

Euphroniaceae]]] Caryocaraceae

N-FIXING CLADE

Fabales

Quillajaceae [Fabaceae [Polygalaceae + Surianaceae]]

Rosales

Rosaceae [[Barbeyaceae [Elaeagnaceae Dirachmaceae Rhamnaceae]] [Ulmaceae

[Cannabaceae [Moraceae + Urticaceae]]]]

Cucurbitales

Anisophylleaceae [[Corynocarpaceae + Coriariaceae] [Cucurbitaceae [Tetramelaceae

[Datiscaceae + Begoniaceae]]]] Apodanthaceae

Fagales

Nothofagaceae [Fagaceae [[Myricaceae + Juglandaceae] [Casuarinaceae

[Ticodendraceae + Betulaceae]]]]

MALVIDROSID II

pgtGeraniales

Geraniaceae [[Melianthaceae + Francoaceae] [Vivianaceae + Ledocarpaceae]]

Myrtales

Combretaceae [[Onagraceae + Lythraceae] [[Vochysiaceae + Myrtaceae]

[Melastomataceae [Crypteroniaceae [Alzateaceae + Penaeaceae]]]]]

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

36

Crossosomatales

[Staphyleaceae [Guamatelaceae [Crossosomataceae + Stachyuraceae]]] [Aphloiaceae

[Geissolomataceae + Strasburgeriaceae]]

Picramniales

Picramniaceae

Sapindales

Biebersteiniaceae Nitrariaceae [Kirkiaceae [Anacardiaceae + Burseraceae]]

[Sapindaceae [Simaroubaceae Rutaceae Meliaceae]]

Huerteales

Gerradinaceae [Dipentodontaceae + Tapisciaceae]

Malvales

Neuradaceae [Thymelaeaceae [Sphaerosepalaceae Bixaceae [Cistaceae

[Sarcolaenaceae + Dipterocarpaceae]] [Cytinaceae + Muntingiaceae] Malvaceae]]

Brassicales

[Akaniaceae + Tropaeolaceae] [[Moringaceae + Caricaceae] [Setchellanthaceae

[Limnanthaceae [[Koeberliniaceae [Bataceae + Salvadoraceae]] [Emblingiaceae

[Pentadiplandraceae [Gyrostemonaceae Resedaceae Stixis and relatives] Tovariaceae

[Capparaceae [Cleomaceae + Brassicaceae]]]]]]]]

Berberidopsidales

Aextoxicaceae + Berberidopsidaceae

Santalales

Erythropalaceae Olacaceae [[Loranthaceae [Misodendraceae + Schoepfiaceae]]

[Opiliaceae + Santalaceae]] Balanophoraceae unplaced

Caryophyllales

[[Droseraceae [Nepenthaceae [Drosophyllaceae [Ancistrocladaceae +

Dioncophylleaceae]]]]] [[Frankeniaceae + Tamaricaceae] [Polygonaceae +

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

37

Plumbaginaceae]]] [Rhabdodendraceae [Simmondsiaceae [[Asteropeiaceae +

Physenaceae] [[Caryophyllaceae [Achatocarpaceae + Amaranthaceae]]

[Stegnospermataceae [Limeaceae [[Lophocarpaceae [Barbeuiaceae [Aizoaceae

[Sarcobataceae Phytolaccaceae Nyctaginaceae]]]] [Molluginaceae [Halophytaceae

Montiaceae Basellaceae Didiereaceae [Talinaceae [Portulacaceae [Anacampseros etc +

Cactaceae]]]]]]]]]]]]]

ASTERIDS

Cornales

[Cornaceae + Nyssaceae] [Hydrangeaceae + Loasaceae] Hydrostachyaceae [Curtisiaceae

+ Grubbiaceae]

Ericales

[Balsaminaceae [Marcgraviaceae + Tetrameristaceae]] [[Polemoniaceae +

Fouquieraceae] Lecythidaceae [[Sladeniaceae + Pentaphylacaceae] [Sapotaceae

[Ebenaceae [Maesaceae [Theophrastaceae [Myrsinaceae + Primulaceae]]]]]

[Mitrastemonaceae Theaceae [Symplocaceae [Styracaceae + Diapensiaceae]]

[[Sarraceniaceae [Roridulaceae + Actinidiaceae] [Clethraceae [Cyrillaceae +

Ericaceae]]]]]]

LAMIIDASTERID I

Unplaced

Metteniusaceae Oncothecaceae Icacinaceae and some unplaced ex-Icacinaceae

Garryales

Garryaceae + Eucommiaceae

Unplaced

Boraginaceae Vahliaceae

Gentianales

Rubiaceae [Gentianaceae [Loganiaceae [Gelsemiaceae + Apocynaceae]]]

Lamiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

38

Plocospermataceae [Oleaceae [Tetrachondraceae [Calceolariaceae

[Gesneriaceae Peltanthera [Pedaliaceae Carlemanniaceae Martyniaceae Stilbaceae

Plantaginaceae Linderniaceae Bignoniaceae Verbenaceae Lamiaceae Paulowniaceae

Schlegeliaceae Thomandersiaceae Phrymaceae Rehmannia Acanthaceae

Scrophulariaceae Orobanchaceae Byblidaceae Lentibulariaceae]]]]]

Solanales

[Montiniaceae [Sphenocleaceae + Hydroleaceae]] [Convolvulaceae + Solanaceae]

CAMPANULIDASTERID II

Aquifoliales

[[Cardiopteridaceae + Stemonuraceae] [Aquifoliaceae [Helwingiaceae +

Phyllonomaceae]]

Asterales

[Rousseaceae + Campanulaceae] Pentaphragmataceae [[Alseuosmiaceae [Phellinaceae

+ Argophyllaceae]] Stylidiaceae [Menyanthaceae [Goodeniaceae [Calyceraceae +

Asteraceae]]]]

Escalloniales

Escalloniaceae

Bruniales

[Bruniaceae + Columelliaceae]

Apiales

Pennantiaceae [Torricelliaceae [Griseliniaceae [Pittosporaceae [Araliaceae

[Myodocarpaceae + Apiaceae]]]]]

Paracryphiales

Paracryphiaceae

Dipsacales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

39

Adoxaceae [Diervillaceae [Caprifoliaceae [Linnaeaceae [Morinaceae [Dipsacaceae +

Valerianaceae]]]]]

LINKS TO ORDERS AND FAMILIES

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MAIN GROUPINGS

Asterids commelinids core eudicots asterid 1 asterid

2 eudicots gymnosperms Magnoliophyta monocots N-fixing claderosids seed plants

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Abietales Acanthales Acerales Acorales Actinidiales Actinostrobales Adoxales Aescul

ales Agavales Aizoales AkanialesAlismatales Alliales Alseuosmiales Alstroemeriales

Altingiales Amaranthales Amaryllidales Amborellales AmbrosialesAmmiales Amomal

es Ancistrocladales Anisophylleales Annonales Anthobolales Apiales Apocynales Apo

nogetonalesAquifoliales Arales Araliales Aralidiales Araucariales Arecales Aristolochi

ales Athrotaxidales Asarales AsclepiadalesAsparagales Asphodelales Asparagales Ast

eliales Atriplicales Aucubales Austrobaileyales Avenales

Balanitales Balanopales Balanophorales Balsaminales Barbeyales Barclayales Batales

Begoniales BerberidalesBerberidopsidales Betulales Biebersteiniales Bignoniales Bix

ales Boraginales Brassicales Brexiales Bromeliales BrunialesBrunoniales Burmannial

es Burserales Butomales Buxales Byblidales

Cactales Callitrichales Calycanthales Calycerales Campanulales Campynematales Can

ellales Cannales CapparalesCaprifoliales Cardiopteridales Carduales Caricales Carle

manniales Caryophyllales Cassiales Casuarinales CelastralesCentrolepidales Cephalot

ales Cephalotaxales Ceratophyllales Cercidiphyllales Chenopodiales Chironiales Chlor

anthalesChrysobalanales Cinchonales Circaeasterales Cistales Citrales Cocosales Col

chicales Columelliales CombretalesCommelinales Connarales Convolvulales Coriarial

es Cornales Corylales Corynocarpales Crassulales CrossosomatalesCucurbitales Cunn

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

40

inghamiales Cunoniales Cupressales Cycadales Cyclanthales Cymodoceales Cynarales

CynomorialesCyperales Cyrillales Cytinales

Daphnales Daphniphyllales Dasypogonales Datiscales Degeneriales Desfontainiales D

iapensiales Didymelales DillenialesDioales Dioncophyllales Dioscoreales Diospyrales

Dipentodontales Dipsacales Droserales

Ebenales Echiales Elaeagnales Elaeocarpales Elatinales Elodeales Emmotales Empetr

ales Ephedrales Ericales EriocaulalesErythropalaless Escalloniales Eucommiales Euph

orbiales Eupomatiales Eupteleales Euryalaless

Fabales Fagales Falcatifoliales Ficales Flacourtiales Flagellariales Fouquieriales Franc

oales Frangulales

Galiales Garryales Geissolomatales Gentianales Geraniales Gesneriales Ginkgoales G

laucidiales Globulariales GnetalesGoodeniales Greyiales Griseliniales Grossulariales

Grubbiales Gunnerales Gyrocarpales Gyrostemonales

Haemodorales Haloragales Hamamelidales Hanguanales Heisteriales Helleborales He

lwingiales Himantandrales HippuridalesHomaliales Hortensiales Huales Huerteales

Hydatellales Hydnorales Hydrangeales Hydrastidales HydrocharitalesHydropeltidales

Hydrostachyales Hypericales Hypoxidales

Icacinales Illiciales Iridales Irvingales Iteales Ixiales

Jasminales Juglandales Julianiales Juncaginales Juncales

Lacistematales Lactoridales Lamiales Lardizabalales Laurales Lecythidales Ledocarpal

es Leitneriales LentibularialesLigustrales Liliales Limnanthales Linales Liriales Loasal

es Lobeliales Loganiales Lonicerales Loranthales LowialesLythrales

Magnoliales Malpighiales Malvales Marathrales Marcgraviales Mayacales Medusagy

nales Medusandrales MelanthialesMelastomatales Meliales Melianthales Meliosmal

es Menispermales Menyanthales Metteniusales MitrastemonalesMiyoshiales Moni

miales Moringales Musales Myricales Myristicales Myrothamnales Myrsinales Myrta

les

Najadales Nandinales Narcissales Nartheciales Nelumbonales Nepenthales Neuradal

es Nitrariales Nolanales NothofagalesNyctaginales Nymphaeales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

41

Ochnales Oenotherales Olacales Oleales Onagrales Oncothecales Opuntiales Orchid

ales Oxalidales

Saxifragales Pandales Pandanales Papaverales Paracryphiales Parasitaxales Paridales

Parnassiales PassifloralesPenaeales Pennantiales Pentaphragmatales Peridiscales P

etiveriales Petrosaviales Phellinales Philydrales PhyllanthalesPhysenales Phytolaccal

es Picramniales Pinales Pinguiculales Piperales Pittosporales Plantaginales Platanale

sPlumbaginales Poales Podocarpales Podophyllales Podostemales Polemoniales Pol

ygalales Polygonales PontederialesPortulacales Posidoniales Potamogetonales Primu

lales Proteales

Quercales Quillajales Quintiniales

Rafflesiales Ranunculales Rapateales Resedales Restionales Rhabdodendrales Rhamn

ales Rhinanthales RhizophoralesRhodorales Rhoipteleales Roridulales Rosales Rouss

eales Rubiales Ruppiales Rutales

Sabiales Salicales Salvadorales Samolales Samydales Sanguisorbales Santalales Sapin

dales Sapotales SarracenialesSaxegotheales Saxifragales Scheuchzeriales Sciadopitya

les Scleranthales Scrophulariales Scyphostegiales SedalesSilenales Simmondsiales S

milacales Solanales Sphenocleales Sphenostemonales Stangeriales Stellariales Stemo

nalesStilbales Stylidiales Styracales Surianales

Taccales Tamales Tamaricales Taxales Taxodiales Tecophilaeales Terebinthales Tern

stroemiales Theales TheligonalesThymelaeales Tiliales Tofieldiales Torricelliales Tov

ariales Tribelales Trilliales Trimeniales Triuridales TrochodendralesTropaeolales Turn

erales Typhales

Ulmales Urticales

Vacciniales Vahliales Vallisneriales Velloziales Veratrales Verbenales Viburnales Vinc

ales Violales Viscales VitalesVochysiales

Welwitschiales Winterales

Xanthorrhoeales Ximeniales Xyridales

Zamiales Zingiberales Zosterales Zygophyllales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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ALPHABETICAL LISTING OF ALL FAMILY NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | K | L | M | N | O | P | Q | R | S | T | U | V | W | X | Y

|Z

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In some cases clicking on a family synonym will send you directly to the subfamily in

which the genus of the basionym is to be found

Abietaceae Abolbodaceae Abrophyllaceae Acalyphaceae Acanthaceae [Acanthochla

mydaceae] Acarnaceaee AceraceaeAchariaceae Achatocarpaceae Achradaceae Achy

ranthaceae Acmopylaceae Acoraceae Acristaceae ActaeaceaeActinidiaceae Actinota

ceae Amentotaxaceae Adenogrammaceae Adoxaceae Aegialitidaceae Aegicerataceae

Aegilopaceae Aeginetiaceaee Aegiphilaceae Aesculaceae Aextoxicaceae Agapanthac

eae Agavaceae AgdestidaceaeAgialidaceae Agrimoniaceae Agrostidaceae Ailanthace

ae Aitoniaceae Aizoaceae Akaniaceae Alangiaceae AlchemillaceaeAldrovandaceae A

lismataceae Alliaceae Allioniaceae Allophylaceae Aloaceae Alopecuraceae Alpiniacea

e AlseuosmiaceaeAlsinaceae Alsodeiaceae Alstroemeriaceae Altingiaceae Alzateacea

e Amaranthaceae Amaryllidaceae Amborellaceae Ambrosiaceae Amentotaxaceae A

mmanniaceae Ammiaceae Amomaceae Ampelidopsaceae Amygdalaceae Amyridacea

eAnacardiaceae Anagallidaceae Anarthriaceae Anchusaceae Ancistrocladaceae Andr

omedaceae AndropogonaceaeAndrostachyaceae Androsynaceae Anemarrhenaceae

Anemonaceae Angelicaceae Anisophylleaceae AnnonaceaeAnomochloaceae Anopter

aceae Anrederaceae Anthemidaceae Anthericaceae Anthobolaceae Antidesmataceae

Antirrhinaceae Antoniaceae Aparinaceae Aphanopetalaceae Aphloiaceae Aphyllanth

aceae Apiaceae ApocynaceaeApodanthaceae Aponogetonaceae Aporusaceae Apose

ridaceae Apostasiaceae Aptandraceae AquifoliaceaeAquilariaceae Aquilegiaceae A

raceae Aragoaceae Araliaceae Aralidiaceaee Araucariaceae Arbutaceae Arceuthidace

aeArceuthobiaceae Arctostaphylaceae Arctotidaceae Ardisiaceae Arecaceae Argoph

yllaceae Arisaraceae AristolochiaceaeAristoteliaceae Arjonaceae Armeriaceae Artem

isiaceae Arthrotaxidaceae Artocarpaceae Arundinaceae ArundinellaceaeAsaraceae A

sclepiadaceae Ascyraceaee Asiraceae Asparagaceae Asperulaceae Asphodelaceae As

pidistraceaeeAsteliaceae Asteraceae Asteranthaceae Asteropeiaceae Astragalaceae

Astrocarpaceae AthanasiaceaeAtherospermataceae Atriplicaceae Atropaceae Aucuba

ceae Aurantiaceae Austrobaileyaceae Austrotaxaceae AvenaceaeAverrhoaceae Avet

raceae Avicenniaceae Azaleaceae Azimaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Balanitaceae Balanopaceae Balanophoraceae Balsameaceae Balsaminaceae Bambusa

ceae Barbaceniaceae BarbeuiaceaeBarbeyaceae Barclayaceae Barringtoniaceae Base

llaceae Bataceae Baueraceae Bauhiniaceae Baxteriaceae BegoniaceaeBehniaceae B

elangeraceae Belloniaceae Bembiciaceae Berberidaceae Berberidopsidaceae Berryac

eaee BersamaceaeBertyaceae Berzeliaceae Besleriaceae Betaceae Betulaceae Biebe

rsteiniaceae Bifariaceae Bignoniaceae BischofiaceaeBixaceae Blakeaceae Blandfordia

ceae Blattiaceae Blepharocaryaceae Blitaceae Blyxaceae Boerlagellaceae Bolivaracea

eBombacaceae Bonnetiaceae Bontiaceae Boopidaceae Boraginaceae Borassaceae B

oroniaceae BoryaceaeBotryodendraceae Bougainvilleaceae Boweniaceae Brassicacea

e Brachycaulaceae Bretschneideraceae BrexiaceaeBromeliaceae Brownlowiaceae Br

unelliaceae Bruniaceae Brunoniaceae Brunsvigiaceae Bryoniaceae BuchneraceaeBuci

daceae Buddlejaceae Buglossaceae Bulbocodiaceae Bumeliaceae Bupleuraceae Burc

hardiaceae BurmanniaceaeBurseraceae Butneriaceae Butomaceae Buxaceae Byblida

ceae Byttneriaceae

Cabombaceae Cacaoaceae Cactaceae Caesalpiniaceae Caladiaceae Calamaceae Calce

olariaceae CalectasiaceaeCalendulaceae Callaceae Callicomaceae Calligonaceae Calli

traceae Callitrichaceae Calochortaceae CalophyllaceaeCalthaceae Calycanthaceae Ca

lyceraceae Cambogiaceae Camelliaceae Campanulaceae CampynemataceaeCanacom

yricaceae Canellaceae Cannabaceae Cannaceae Canopodaceae Canotiaceae Cansjera

ceae CapparaceaeCaprariaceae Caprifoliaceae Cardiopteridaceae Carduaceae Carica

ceae Carissaceae Carlemanniaceae CarpinaceaeCarpodetaceae Cartonemataceae Ca

ryocaraceae Caryophyllaceae Casearieaceae Cassiaceae CassipoureaceaeCassythacea

e Castaneaceae Castelaceae Casuarinaceae Catesbaeaceae Cathedraceae Cecropiace

ae CedraceaeCedrelaceae Celastraceae Celosiaceae Celtidaceae Centaureaceae Cen

trolepidaceae Centroplacaceae CepaceaeCephalanthaceae Cephalotaceae Cephalota

xaceae Cerastiaceae Ceratoniaceae Ceratophyllaceae CerberaceaeCercidiphyllaceae

Cercocarpaceae Cercodiaceae Cereaceae Cerinthaceae Ceroxylaceae Cestraceae Cev

alliaceaeChailletiaceae Chamaedoraceae Chamaemoraceae Chamelauciaceae Chaun

ochitonaceae Chelidoniaceae ChelonaceaeChenopodiaceae Chimonanthaceae Chingi

thamnaceae Chionographidaceae Chiranthodendraceae ChironiaceaeChlaenaceae Ch

loanthaceae Chloranthaceae Chloridaceae Chlorogalaceae Chrysobalanaceae Cicerac

eae CichoriaceaeCimicifugaceae Cinchoniaceae Circaeaceae Circaeasteraceae Cissac

eae Cistaceae Citraceae Clematidaceae CleomaceaeClethraceae Clusiaceae Cneorac

eae Cnestidaceae Cnicaceae Cobaeaceae Cochlospermaceae Cocosaceae Coffeaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Colchicaceae Coleogynaceae Columelliaceae Combretaceae Commelinaceae Comocl

adiaceae Compositae CompsoaceaeConantheraceae Connaraceae Conostylidaceae C

onvallariaceae Convolvulaceae Cordiaceae CoreopsidaceaeCoriandraceae Coriariacea

e Coridaceae Corispermaceae Cornaceae Corokiaceae Coronillaceae Correaceae Cor

rigiolaceaeCorsiaceae Corydalaceae Corylaceae Corynocarpaceae Coryphaceae Cost

aceae Cotyledonaceae CoulaceaeCoutariaceae Coutoubiaceae Crassulaceae Crescen

tiaceae Cressaceae Crinaceae Crocaceae CroomiaceaeCrossosomataceae Crotonacea

e Cruciferae Cryptaceae Crypteroniaceae Cryptocorynaceae CryptomeriaceaeCtenol

ophonaceae Cucurbitaceae Cunninghamiaceae Cunoniaceae Cupressaceae Curcumac

eae Curtisiaceae CuscutaceaeCyananthaceae Cyanastraceae Cynanchaceae Cyanella

ceae Cycadaceae Cyclanthaceae CyclantheraceaeCyclocheilaceae Cydoniaceae Cymo

doceaceae Cynaraceae Cynocrambaceae Cynomoriaceae Cyperaceae CyphiaceaeCyp

hocarpaceae Cypripediaceae Cyrillaceae Cyrtandraceae Cyrtanthaceae Cytinaceae

Dacrycarpaceae Dactylanthaceae Damasoniaceae Daphnaceae Daphniphyllaceae Das

ypogonaceae DatiscaceaeDaturaceae Daucaceae Davidiaceae Davidsoniaceae Decais

neaceae Deeringiaceae Degeneriaceae DelphiniaceaeDendrophthoaceae Columelliac

eae Detariaceae Dialypetalanthaceae Dianellaceae Dianthaceae DiapensiaceaeDicha

petalaceae Dichondraceae Diclidantheraceae Dicrastylidiaceae Dictamnaceae Didiere

aceae DidymelaceaeDidymocarpaceae Diegodendraceae Diervillaceae Digitalidaceae

e Dilatridaceae Dilleniaceae Dioaceae DionaeaceaeDioncophyllaceae Dioscoreaceae

Diosmaceae Diospyraceae Dipentodontaceae Diphylleiaceae Diplolaenaceae Dipsaca

ceaeDipterocarpaceae Dirachmaceae Disanthaceae Diselmaceae Dodonaeaceae Do

mbeyaceae Donatiaceae DorsteniaceaeDortmannaceae Doryanthaceae Drabaceae D

racaenaceae Dracontiaceae Drimyidaceae Droseraceae DrosophyllaceaeDryadaceae

Duabangaceae Duckeodendraceae Dulongiaceae Durantaceae Durionaceae Dysphani

aceae

Ebenaceae Eccremidaceae Ecdeiocoleaceae Echiaceae Echinopaceae Ehretiaceae Ela

eagnaceae ElaeocarpaceaeElegiaceae Elatinaceae Ellisiaceae Ellisiophyllaceae Elodea

ceae Elytranthaceae Embeliaceae Emblingiaceae EmmotaceaeEmpetraceae Encepha

lartaceae Engelhardtiaceae Enhalaceae Epacridaceae Ephedraceae Ephemeranthacea

eEpilobiaceae Epimediaceae Eragrostidaceae Eremolepidaceae Eremosynaceae Erica

ceae Erinaceae EriocaulaceaeEriogonaceae Eriospermaceae Erodiaceae Erycibaceae

Eryngiaceae Erysimaceae Erythroniaceae ErythropalaceaeErythrospermaceae Erythro

xylaceae Escalloniaceae Eschscholziaceae Eucomidaceae Eucommiaceae Eucryphiace

aeEuonymaceae Eupatoriaceae Euphorbiaceae Euphrasiaceae Euphroniaceae Eupo

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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matiaceae Eupteleaceae EuryalaceaeEustrephaceae Euthemidaceae Eutocaceae Exb

ucklandiaceae Exocarpaceae

Fabaceae Fagaceae Falcatifoliaceae Ferulaceae Festucaceae Ficaceae Ficoidaceae Fit

zroyaceae FlacourtiaceaeFlagellariaceae Flindersiaceae Foetidiaceae Forestieraceae

Fothergillaceae Fouquieriaceae Fragariaceae FrancoaceaeFrangulaceae Frankeniacea

e Fraxinaceae Fraxinellaceae Freycinetiaceae Fritillariaceae Fuchsiaceae Fumariacea

eFunkiaceae

Gaiadendraceae Galacaceae Galanthaceae Galaxiaceae Galedupaceae Galeniaceae G

aliaceae Garciniaceae GardeniaceaeGardneriaceae Garryaceae Geissolomataceae Ge

itonoplesiaceae Gelsemiaceae Geniostomaceae GentianaceaeGeonomataceae Geosir

idaceae Geraniaceae Gerrardinaceae Gesneriaceae Gethyllidaceae Gilliesiaceae Gina

lloaceaeGinkgoaceae Gisekiaceae Gladiolaceae Glaucidiaceae Glechomaceae Glinace

ae Globulariaceae Gnaphaliaceae GnetaceaeGoetzeaceae Gomortegaceae Gomphiac

eae Gomphrenaceae Gonystylaceae Goodeniaceae Gordoniaceae GouaniaceaeGoup

iaceae Gramineae Gratiolaceae Grewiaceae Greyiaceae Grielaceae Griseliniaceae Gr

onoviaceae GrossulariaceaeGrubbiaceae Guaiacanaceae Guamatelaceae Guettardac

eae Gunneraceae Gustaviaceae Guttiferae GyrocarpaceaeGyrostemonaceae

Hachetteaceae Haemanthaceae Haemodoraceae Halesiaceae Hallieraceae Halocarpa

ceae Halophilaceae HalophytaceaeHaloragaceae Hamamelidaceae Hanguanaceae H

aptanthaceae Harmandaceae Hebenstretiaceae HectorellaceaeHederaceae Hedyosm

aceae Hedyotidaceae Hedysaraceae Heisteriaceae Heleniaceae Heliamphoraceae H

elianthaceaeHelianthemaceae Helichrysaceae Heliconiaceae Helicteraceae Heliotropi

aceae Helleboraceae Heloniadaceae HelosidaceaeHelwingiaceae Hemerocallidaceae

Hemimeridaceae Hemiphylacaceae Henriqueziaceae Henslowiaceae Hermanniaceae

Hernandiaceae Herniariaceae Herreriaceae Hesperocallidaceae Heterantheraceae He

teropyxidaceae HeterostylaceaeHibbertiaceae Hibiscaceae Hilleriaceae Himantandra

ceae Hippocastanaceae Hippocrateaceae HippomanaceaeHippophaeaceae Hippurida

ceae Hirtellaceae Holacanthaceae Homaliaceae Hopkinsiaceae Hoplestigmataceae H

ordeaceaeHornschuchiaceae Hortensiaceae Hortoniaceae Hostaceae Houstoniaceae

Huaceae Huerteaceae HugoniaceaeHumbertiaceae Humiriaceae Hyacinthaceae Hyd

atellaceae Hydnoraceae Hydrangeaceae Hydrastidaceae HydrillaceaeHydrocharitacea

e Hydrocotylaceae Hydrogetonaceae Hydroleaceae Hydropeltidaceae Hydrophylacac

eae HydrophyllaceaeHydrostachyaceae Hymenocardiaceae Hypecoaceae Hyperanth

eraceae Hypericaceae Hypopityaceae HypoxidaceaeHypseocharitaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta

ceae ImperatoriaceaeInocarpaceae Inulaceae Ionidiaceae Iriarteaceae Iridaceae Irvi

ngiaceae Isnardiaceae Isophysidaceae IteaceaeStrasburgeriaceae Ixiaceae Ixioliriace

ae Ixonanthaceae

Jalapaceae Jamboliferaceae Japonoliriaceae Jasionaceae Jasminiaceae Johnsoniaceae

Joinvilleaceae JuglandaceaeJulianiaceae Juncaceae Juncaginaceae Juniperaceae Jus

siaeaceae Justiciaceae

Kadsuraceae Kaliphoraceae Kalmiaceae Kaniaceae Kiggelariaceae Kingdoniaceae Kin

giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae

Krameriaceae

Labiatae Lacandoniaceae Lachenaliaceae Lacistemataceae Lactoridaceae Lactucaceae

Lagerstroemiaceaee LamiaceaeLampsanaceae Lanariaceae Langsdorffiaceae Lantan

aceae Lapageriaceae Lardizabalaceae Lasiopetalaceae LauraceaeLawsoniaceae Laxm

anniaceae Lecythidaceae Ledaceae Ledocarpaceae Leeaceae Leguminosae Leitneriac

eae LemnaceaeLennoaceae Lentibulariaceae Lentiscaceae Leoniaceae Leonticaceae

Lepidobotryaceae LepidocarpaceaeLepidocaryaceae Lepidocerataceae Leptaulaceae

Leptospermaceae Lepturaceae Lepuropetalaceae LeuctenbergiaceaeLibocedraceae

Licaniaceae Ligustraceae Lilaeaceae Liliaceae Limeaceae Limnanthaceae Limnocharit

aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace

ae Linderniaceae Lindleyaceae Linnaeaceae LippayaceaeLiriaceae Liriodendraceae Li

ssocarpaceae Littorellaceae Loasaceae Lobeliaceaee Loganiaceae LomandraceaeLoni

ceraceae Lopeziaceae Lophiocarpaceae Lophiolaceae Lophiraceae Lophophytaceae L

ophopyxidaceae LoranthaceaeLotaceae Lowiaceae Lupulaceaee Luxemburgiaceae Lu

zuriagaceae Lychnidaceae Lyciaceae LyginiaceaeLygodisodeaceae Lysimachiaceae Ly

thraceae

Macarisiaceae Mackinlayaceae Madiaceae Maesaceae Magnoliaceae Malaceae Male

sherbiaceae MalortieaceaeMalpighiaceae Malvaceae Mangiaceae Manicariaceae Ma

paniaceae Marantaceae Marathraceae MarcgraviaceaeMartyniaceae Mastixiaceae

Matricariaceae Maundiaceae Mayacaceae Medeolaceae Medusagynaceae Peridiscac

eaeMelaleucaceae Melampyraceae Melanophyllaceae Melanthiaceae Melastomatac

eae Meliaceae Melianthaceae MelicaceaeMeliosmaceae Melittidaceae Melochiacea

e Memecylaceae Mendonciaceae Menispermaceae Menthaceae MenyanthaceaeMe

nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

47

Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

Nypaceae Nyssaceae

Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

ceae Papilionaceae Pappophoraceae PapyraceaeParacryphiaceae Parasitaxaceae Par

ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

48

hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

49

cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

50

ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 12: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

12

a family or a subfamily That way surely lies madness and worse the discredit of our

discipline

There are similar issues whatever naming system is used Thus in phylogenetic

naming (Baum et al 1998 for an example but cf Baum et al 2004 for the PhyloCode

see Cantino amp De Queiroz 2006) an unflagged hierarchy is used in which any

terminations of names used are uninformative about the relative position of taxa If one

adopts the principle of phylogenetic naming one indeed does not have to worry about

the nomenclatural consequences caused by lumping or splitting any well-supported

clade can be named without affecting the name of more or less inclusive clades

Unfortunately unflagged hierarchies have very serious deficiences as communication

devices because they lack one aspect essential in language biological or otherwise -

they contain no intrinsic information about the relationships of the group in question to

others (eg Pfeil amp Crisp 2005 Stevens 2006a) Recent suggestions for using prefixes like

Apo- and Pan- to PhyloCode names will however allow limited information of this

kind to be conveyed but only as it pertains to individual branches and current

proposals do not even mandate that the prefixes be employed consistently In any

event such proposals simply prevent the potential tripling of the number of quite

different names used to describe different aspects of a phylogenetic tree over those

used to name monophyletic groups pure and simple In general where n is the number

of extant species in a group the number of clades in such a group = n-1 (Species will

also need names too for their names see Dayrat et al 2008) Importantly here too

consensus over the clade names commonly learned by students and used in herbaria is

needed otherwise communication will be impeded the names themselves will provide

no guidelines as to which should be chosen The situation is of course more complicated

than this Terminations that convey ideas of rank in a phylogenetic classification can also

be used in phylocode names - however there they will carry no implications of rank

How they will be used is another matter of course

Of course there are other philosophies of classification and some still prefer

evolutionary classifications There classificatory principles differ substantially from those

followed here eg the recognition of paraphyletic taxa may be permitted however

detailed reasons for prefering the taxa that are recognised are rarely given although

nature and natural groups are often mentioned (cf Stuessy amp Koumlnig 2008) For

summaries of commonly used systems see Brummitt (1992) and Mabberley (2008)

new evolutionary systems appear every year or so However even those who allow or

promote the recognition of paraphyletic groups (eg Grant 2003 Thorne 2007

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

13

Heywood et al 2007) may find it of interest to examine a system recognizing only

monophyletic groups it provides a rather different understanding of evolution

Evolutionary classifications in general try and combine phylogeny and

morphological gaps although that is no easy thing to do - it is akin to combining chalk

and cheese (for an attempt to make this impossible task seem to be more objective see

Stuessy amp Koumlnig 2008) However there is a principle from evolutionary classification

that is relevant and I think quite useful here The size of the gap between two groups

tends to be inversely proportional to the sizes of the groups involved (Davis amp Heywood

1963) One can imagine a situation in which a large group is formally divided even

although the distinguishing characters of the two are weak whereas a smaller group

similarly divisable will be left intact

To summarize If hypotheses of phylogeny remain stable we should be able to base

a stable classification on that phylogeny and then get on with our work that is testing

the phylogenies we have elucidating phylogenies in areas where relationships are

unclear studying the evolution of morphology describing species etc In this context

the spread of the Angiosperm Phylogeny Group system (see below) and its widespread

utilisation in technical literature also floras (eg van der Meijden 2005) textbooks (eg

Simpson 2006 Judd et al 2007 [third edition]) dictionaries (Mabberley 2008) more

popular literature (eg Souza amp Lorenzi 2005 Spears 2006) and as an outline for a new

herbarium sequence (Haston et al 2007) is gratifying The posibility that one might be

able to develop a stable phylogeny-based classification of families and in particular

orders represents a dramatic turn-around from the pessimistic attitude about such

higher-level groupings expressed by Davis and Heywood (1963) and Thorne (1976) the

latter even suggesting that we should bury forever the metaphor of the phylogenetic

tree as highly unrealistic (ibid p 56) Returning to Godfray and Knapps (2004) users

of classifications who want a stable informative and accessible classification that

enables easy identification - unfortunately they want cake with everything and cannot

get it - these pages attempt to satisfy as many of their needs as possible but phylogeny

and monophyly are the primary shapers of the APG classification

On this classification in particular

Here I very largely follow the Angiosperm Phylogeny Group classification (APG

2003) Any differences are not to be interpreted as differences in principle simply that

new phylogenies continue to be published and here I attempt to provide an overview

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

14

of current ideas of higher-level relationships of seed plants The Angiosperm Phylogeny

Group classification is based on well-supported relationships evident in the numerous

molecular studies that began to appear in the late 1980s (see APG 1999 for the

principles underlying the classification) but recent phylogenetic work does not

contradict the major outlines of the trees used by APG II (2003) or even those in APG

I (APG 1999) The APG classification has for the most part been conservative and I

am similarly conservative here For the names and authorities I follow APG III (2009)

although the names of the authors may not always be correct these being in something

of a state of flux for the superordinal names used see Chase and Reveal (2009)

although classification at this level here is not exhaustive - only clades with several

orders have superordinal names

Some changes in our ideas of relationships and hence in the clades we talk about

are particularly likely in parts of Lamiales Caryophyllales and Malpighiales for example

although this will not affect the orders themselves Thus some changes to clade

circumscriptions are to be expected although I do not expect them to be substantial

but changes are neither a defect of cladistics nor a necessary consequence of the use of

molecular data Clades are hypotheses of relationships and as hypotheses they may be

overturned However work since APG I has not suggested other than minor changes

in the compositions of the orders even if the odd genus or even family is turning out to

be seriously misplaced - recent examples are Hydatellaceae (from monocots-Poales to

Nymphaeales Saarela et al 2007) Guamatelaceae (from Rosales-Rosaceae to

Crossosomatales Oh amp Potter 2006) and Perrottetia and Bhesa (from Celastrales-

Celastraceae to Huerteales and Malpighiales respectively Zhang amp Simmons 2006) The

main changes have been clarification of the relationhips of individual families or groups

of families that were of uncertain position eg of Chloranthaceae (Moore et al 2007)

Ceratophyllaceae (Jansen et al 2007) and unplaced asterid II families (Winkworth et al

2008a) As mentioned above phylogenies in the area of main interest in these pages are

overwhelmingly tree-like Although there is evidence for acquisition of host plant genes

by parasites (see below) even wider but inexplicable transfer of mitochochondrial

genes in plants like Amborella (Bergthorsson et al 2004 cf Goremykin et al 2009a)

and even transfers of nuclear genes (Vallenback et al 2008) these are the exception

rather than the rule

In cases where the Angiosperm Phylogeny Group II allowed alternatives as to the

limits of families - Papaveraceae in the broad sense or Papaveraceae plus

Pteridophyllaceae plus Fumariaceae Proteaceae in the broad sense or Proteaceae plus

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

15

Platanaceae - the choices made here follow common usage eg as in textbooks like

Judd et al (2007) and Simpson (2006) and particularly in the new edition of

Mabberleys The Plant Book (Mabberley 2008) This latter is itself an attempt to reflect a

consensus the result of taking the opinions of botanists at several meetings A largely

similar consensus is reflected in the APG III classification (APG 2009) where you will

see that alternative classifications have been dispensed with For many people the

existence of such alternative classifications will simply confuse so agreement over

which groupings to use when alternatives are permitted and then dispensing with the

alternatives that are not commonly used seems reasonable I am making the few

changes that are needed to bring the classification here in line with these others but

only gradually since there is not enough time in each day

As already mentioned higher-level relationships in general and the composition of

orders in particular have always presented something of a challenge to systematists

(eg Davis amp Heywood 1963) That being said the composition of some clades like

Apiales Crossosomatales and Pandanales is decidedly unexpected However it is

interesting to see that these higher level clades are generally accepted even in works

with different classificatory philosophies (eg Heywood et al 2007) For clades like

Malpighiales many of the family groupings within Asparagales etc attempts to find

distinctive characters have largely failed (but see Endress amp Matthews 2006a also

the Apomorphies page here) Interestingly as with families some groupings suggested

by molecular studies are supported by morphological andor chemical characters that

have long been known sometimes for over a hundred years the relationship between

Pittosporaceae and ApiaceaeAraliaceae are a case in point (Hegnauer 1969b and

references) As our knowledge of morphology and chemistry improves we can hope for

improvements in the characterisations of clades at all levels

In many cases the new family limits of the Angiosperm Phylogeny Group (see

APG 1999 2003) are not really controversial although changes from the limits

commonly accepted only a decade ago are sometimes quite dramatic (eg Wagenitz

1997) Thus the split of the old Saxifragaceae sl is necessitated by its extreme

polyphyly as also with Icacinaceaesl and Cornaceae sl However the limits and

relationships of such groups had long been considered to be unsatisfactory but until

recently there had been no real reason to prefer one proposed arrangement over

another It is generally accepted that the limits of Lamiaceae and Verbenaceae have to

be redrawn although there is as yet no compelling evidence that the redrawn taxa are

not sister taxa (there is no evidence that they are) But whatever their relationships the

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

16

content of the clades has changed considerably and incidentally they are now easier to

identify than before the decision to recognise two families is not difficult The same is

true for Salicaceae and Achariaceae (Malpighiales) two previously small families that

have received the bulk of the old Flacourtiaceae Clade and hence taxon limits remain

difficult around Theaceae (Ericalessee alsoSladeniaceae Pentaphylacaceae)

and Euphorbiaceae (Malpighiales see

also Phyllanthaceae Picrodendraceae Putranjivaceae etc) Here current groupings

may not represent quite such dramatic changes in our understanding of relationships

Indeed parts of the old Euphorbiaceae that were separated may yet go back together

particularly Phyllanthaceae and Picrodendraceae (see Wurdack et al 2004 Davis et al

2005) even if Theaceae should indeed be dismembered (Geuten et al 2004) However

given that there is no molecular evidence that currently warrants combining all the

segregates of Euphorbiaceae and even if some do come together the clades the

families represent suggest novel groupings not recognised in current classifications

Furthermore Rafflesiaceae appear to be embedded within Euphorbiaceae s str (Davis

et al 2007) so the family is divided Of course maintaining even a moderately narrowly-

circumscribed Euphorbiaceae would entail reducing the iconic Rafflesiaceae to

synonymy Relationships in core Caryophyllales especially

around Phytolaccaceae andNyctaginaceae and also Portulacaceae are incompletely

understood and refashioning of taxon limits will doubtless be needed as cladistic

relationships become apparent (eg see Nyffeler 2007 Ogburn 2007a b) Some

groupings in the old Icacinaceaeand Olacaceae also remain unclear

The discovery of the relationships of parasitic and aquatic groups have presented a

particular challenge to systematists Morphologically some of these plants are so highly

modified that interpretation of the plant body in conventional terms is difficult or even

impossible Thus parasitic groups (broad-leaved angiosperms only) such as Rafflesiaceae

are hard to place since both the vegetative body and the flowers are changed almost

beyond recognition (flowers of rafflesiaceae are a good example of gigantism - Davis et

al 2007) Furthermore plastid gene sequences may be difficult or impossible to obtain

the chloroplast DNA in particular being highly degraded and the rate of molecular

change in general may be high so the problem of long-branch attraction is serious (eg

Duff amp Nickrent 1997 Nickrent et al 1998 Caddick et al 2002a G Petersen et al

2006b) Echlorophyllous sapromycoheterotrophic taxa mostly known from monocots

present similar problems However progress is being made placements for

Apodanthaceae Rafflesiaceae Mitrastemonaceae Cytinaceae and Cynomoriaceae have

recently been suggested (Barkman et al 2004 2007 Davis amp Wurdack 2004 Nickrent et

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

17

al 2004 Davis et al 2007) it seems likely that Burmanniaceae sl are polyphyletic but

both parts are in Dioscoreales (Merckx et al 2006) and relationships within the largely

hemiparasitic Santalales are also gradually being clarified (Maleacutecot 2002) The inclusion

of parasitic taxa in general molecular analyses can cause conniptions (eg Nickrent et al

2004 Davis et al 2004 Chase et al 2006 G Petersen et al 2006b) and there can be

horizontal transmission of genes (eg Davis amp Wurdack 2005 Vitaceae to Rafflesiaceae

Barkman et al 2007 the mitochondrial atp1 gene commonly moves)

Similarly in water plants neither vessels in particular nor much xylem in general is

needed leaves are highly modified and water-mediated pollination if adopted may

well be associated with major changes in floral morphology Here too recent molecular

studies suggest that aquatic groups with hitherto problematic relationships may find

homes Thus Podostemaceae are close to Clusiaceae and Calophyllaceae and are sister

to Hypericaceae (Malpighiales Kita amp Kato 2001) Hydatellaceae which used to be in

Poales are part of Nymphaeales (Saarela et al 2007) and Hydrostachyaceae may be

close to Hydrangeaceae (Cornales Xiang et al 2002) or perhaps should be placed in

Lamiales note that in the first two cases in particular there are morphological and

chemical features that support such a move If Podostemaceae are indeed close to

Clusiaceae I look forward to seeing hypotheses to explain how the dramatic changes in

the vegetative body that have made Podostemaceae so problematic for generations of

systematists took place That conventional wisdom has trouble in understanding or

explaining how the morphologies of groups like Clusiaceae and Podostemaceae can be

related is largely a problem with conventional wisdom Ceratophyllaceae seem to be

finding a position as sister to eudicots (see Moore et al 2007) but their morphology is

so derived that there is no morphological evidence of which I am aware for this

relationship There are also similar although less extreme morphological problems with

many plants pollinated by wind

In such situations especially with some water plants mycoheterotrophs and

parasites a variety of characters may be affected and particularly under such

circumstances groups have tended to be formed using a few or even only a single

character that seems to provide evidence of relationships Add to this the tendency to

weight some characters particularly strongly on a priori grounds and the result was

either the recognition of conglomerate taxa such as Amentiferae which now appear to

be highly polyphyletic or the segregation of families like Plantaginaceae s str (now

much expanded) and Leitneriaceae (now in Simaroubaceae) - all of these are more or

less wind pollinated

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

18

Even although relationships of many parasitic and aquatic groups are being

clarified and at least some of the proposed phylogenetic positions seem to be well

supported it is indeed often still very difficult to understand how such groups have

evolved from their more morphologically conventional relatives However we have had

all sorts of preconceptions as to how characters should evolve Thus we used to

assume that features like highly scalariform vessel perforation plates or the complete

absence of vessels or a flower with an androecium that had many stamens a superior

ovary or separate petals were necessarily plesiomorphic or primitive and conversely

a simple perforation plate and the presence of vessels an androecium with few

stamens inferior ovary or petals that were connnate were almost necessarily

apomorphic or advanced (but cf eg Stebbins 1951) Such assumptions are incorrect

(eg Soltis et al 2005b) Carpels may become secondarily free carpels may fail to close

the seeds then developing outside the confines of the carpel as in some Aspagaraceae-

Nolinoideae Violaceae Berberidaceae Malvaceae-Sterculioideae etc) in Peliosanthes

teta perhaps the only species in Peliosanthes(Asparagaceae-Nolinoideae) the ovary

varies from superior to inferior (Jessop 1976 species limits here need close

investigation see also Kuzoff et al 2001 and Soltis amp Hufford 2002 Saxifragaceae

Apiales etc) many-seeded carpels can evolve from few-seeded carpels

(Razafimandimbison et al 2008) monoecy may be derived from dioecy (Schaefer amp

Renner 2010 and references) Classic studies such as those by Babcock (eg 1947)

on Crepis that assumed that evolution - in this case of the karyotype in particular - was

unidirectional have needed comprehensive re-evaluation (Enke amp Gemeinholzer 2008)

Most if not all characters have reversed andor evolved in parallel as is clear in the

discussion of ovary position here especially in Poales Saxifragales and Asparagales and

also in comments on such morphologically distinctive taxa (when compared with their

immediate relatives) such as Menyanthaceae and especially Pittosporaceae Parallel

evolution may occur even at the level of amino acid substitution as in the independent

acquisition of the phosphoenolpyruvate carboxylase (pepC) gene in C4 photosynthesis in

grasses (Christin et al 2007b see also Blaumlsing et al 2000)

Thinking of how such characters have been used in the past in classifications at the

generic level the results have been similar to those just discussed above for parasitic

plants and others Relying too much on animal pollination syndromes to mark generic

boundaries has all too often led to taxa that are highly unsatisfactory phylogenetically

(see eg Acanthaceae Bignoniaceae Campanulaceae Ericaceae Lamiaceae

Orchidaceae) and over-reliance on characters of fruit and seed (see eg Brassicaceae

and Apiaceae) has also led to unsatisfactory generic limits Again the more general

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

19

problem is the use of one or a very few characters that have been weighted a priori to

structure classifications (see also Garciacutea et al 2009) although it is perhaps not

surprising that such an approach sometimes results in the delineation of taxa that have

indeed turned out to be quite distinct (cf eg Burtt Astragalus versus Oxytropis)

Taxa recognized here are monophyletic yet many are polythetic at the

morphological level that is they lack unique features characterizing (defining) all and

only members of that family they can be recognised phenetically only by the unique

combinations of characters that they posess This is the result of evolution any

synapomorphy characterizing a taxon may be lost or modified beyond easy recognition

in some of its members or the synapomorphy may appear to be identical to a feature

that has evolved in parallel in a quite unrelated plant That plant groups are polythetic is

almost as much a feature of monophyletic taxa as the paraphyletic taxa common in

evolutionary classifications Some families now include substantial variation as

phenetically distinct derived groups are placed in their proper phylogenetic position -

examples are Ericaceae which include the erstwhile Empetraceae a wind-pollinated

group that is florally very different and the overwhelmingly large-flowered and animal-

pollinated Plantaginaceae which now include several derived small-flowered aquatic

and wind-pollinated groups that were previously placed in separate families

Thus some families as delimited here may not be easy to recognize However

remember that detecting relationships - use whatever characters you can even if they

are not obvious - and naming a plant - focus on easy-to-see characters that may not

reflect relationships - are quite different problems Taxa although natural may not be

readily recognizable indeed it was in exactly this context that Lamarck worked out the

basic principles of writing dichotomous keys in 1778 (see Scharf 2007 for keys and the

like) Of course Lamarcks idea of nature was very different from ours - he thought that

there was some kind of continuum of form on which living organisms were to be

situated with no real gaps anywhere - but this meant that his genera (for example)

might well not be sharply distinct from each other even if each were part of the real

continuum that was life Lamarcks separation of the establishment of relationships and

groupings in a system and identification of the plants in that system have been widely

accepted (but cf eg Godfray amp Knapp 2004 many of the contributors to Heywood et

al 2007) Perhaps the best way of identifying plants at the family level is by well-made

multiple access keys as in Watson and Dallwitz (1992a onwards family limits there may

differ substantially from those adopted here) Multiple access keys free users from the

constraints of dichotomous keys in which particular characters are needed at each step

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

20

of the identification process before there can be further progress Instead those

characters that happen to be evident on a specimen can be used in whatever order is

convenient when linked to illustrations glossaries etc their power is enormous (see

Dallwitz et al 2000 [2006] for the principles underlying their construction and use)

Nevertheless dichotomous keys such as those of Hutchinson (1973) and Franz Thonner

(Geesink et al 1981) have their uses Of course taxa coming out adjacent in keys may

well not be at all related

(When identifying large numbers of plants even more efficient than either style of

identification and certainly lots more fun is sight identification Unless you have a

photographic memory you have to build up your knowledge of comparative plant

morphology - on which the ability to make accurate identifications depends - by

repeated observation When faced with an unknown plant I always look for leaf teeth

and stipules distinctive hair types especially stellate and T-shaped hairs glands and

punctations of any sort the presence of latex or other exudate and check leaf insertion

smelling crushed leaves can also be helpful In this context nodal anatomy can usually

be checked using a razor and a hand lens or even simply carefully examining leaf scars

The short paragraphs added after most families may help in confirming familial

identifications)

For the record and for the little that it is worth there are 4 orders and 13 families

of gymnosperms characterised on these pages and together they include some 82

genera and 947 species For angiosperms comparable figures are 56 orders 445

families 13208 genera and 261750 species (of which monocots include 11 orders 89

families 2759 genera and 52760 species) Note however that higher mathematics

was never my strong point and anyway these are pretty meaningless figures even for

species which many (but not all) might concede smacked slightly more of reality than

other taxa estimates range as high as 422000 (Govaerts 2001) Furthermore numbers

of genera and species change daily Nevertheless as emphasized here families are

useful in teaching we as a community can ensure that their limits remain largely stable

and by concentrating on relatively few of them one can gain some familiarity with much

of the worlds flora For summaries of the sizes of orders and families and of the general

arrangement followed in these pages see also the Statistics page itself

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

21

A PHYLOGENETIC TREE OF SEED PLANTS (THE MAIN TREE)

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

22

The Main Tree above is a conservative summary of well supported relationships

between the major clades of extant seed plants Reading - interpreting and

understanding - such a phylogenetic tree is not necessarily easy so reading a tree may

require a bit of practice OHara (eg 1988 1992) provides an accessible introduction to

thinking and talking about history in the context of looking at phylogenetic trees More

recently Krell and Cranston (2004) Crisp and Cook (2005) and others have emphasized

how careful one must be when interpreting and talking about evolution in ladderized

trees in particular and phylogenetic trees in general Gregory (2008) provides a

particularly detailed discussion of this problem In ladderized trees the smaller (in terms

of numbers of terminals) sister taxon is consistently shown on the same side at every

node thus the trees tend to be pectinate like the teeth of a comb however the

horizontal axis that is as a result so evident has no polarity or particular direction The

use of the adjective basal when discussing phylogenetic trees is especially dangerous

(see also Wojciechowski et al 2004 D Soltis et al 2005b) When I use the term and the

context is not otherwise clear I am referring to the pectinations at the base of a

ladderised tree Remember that when talking about sister taxa one can never be basal

to the other or older or younger although members of one can be more derived - in the

sense of having more apomorphies - than the other (but even then one has to be

careful) The words primitive and advanced and lower and higher should

practically never be used especially when talking about taxa Amborellaceae and

Pinaceae are sister to all other flowering plants and Pinales and are basal to the crown

groups of all other flowering plantsPinales respectively but that does not mean they

are thereby primitive Pinaceae in particular have numerous apomorphies The word

plesiomorphic is far less loaded than primitive and can be used to talk about

individual characters

Most trees here have been more or less ladderized as is the one above The

ladderization may be imperfect for example you can see that on the Main Tree

asterids with ten orders follow rosids and relatives which have seventeen although

this is in part because of the currently unresolved nature of relationships between the

main clades of core eudicots (this is likely to change soon) The Main Tree could have

been drawn with Amborella Acorus or a host of other taxa at the far right without

offending any relationships phylogenetic trees are like mobiles the only fixed points

being the nodes Despite the problems just mentioned pectination interpreted

carefully has its value and it is not an arbitrary process As one reads the terminals of a

pectinate tree from left to right adjacent terminals will be separated by apomorphies

that will cumulate in an appropriate fashion When thinking of a book or a herbarium

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

23

sequence (see Haston et al 2007) this is of some value Nymphaeales and

Austrobaileyales are here adjacent on the tree but they could be separated by

hundreds of families in the sequence merely by rotating only the node from which

Nymphaeales and their sister group (all other angiosperms minus about 75 species)

arise If adjacent in a book or herbarium or as in these summary pages then it is

relatively easy to relate their characters both apomorphic and plesiomorphic but if

separated by hundreds of pages or two floors in a large building then it is less easy to

get anything from the sequence (Since all orders in the pages of the main part of this

site are preceded by the apomorphies of all nodes immediately below them in the seed

plant phylogeny and because of the linkages that have been built in to the site this

problem is much less serious there there is no sequence) So the particular sequence of

all seed plants that is based on a phylogenetic classification but is to be used in herbaria

or any other place where taxa have to be arranged linearly as in this part of APweb

will be one that maximizes the number of taxa that are both successive branches of the

tree and placed successively in the sequence Since specimens are generally filed under

families the outline of a new family sequence for arranging herbaria and books can now

be suggested (Haston et al 2007) and such a standardized sequence will further help

teaching and learning about plants

ON THINKING ABOUT APOMORPHIES

Identifying apomorphies is important because understanding the evolution of

morphology in the broad sense - ie including anatomy chemistry etc - in the context

of diversification is one of our major goals For identifying apomorphies several

preconditions must be met One needs to have an accurate well supported phylogeny

one has to have examined the right taxa both from the point of view of morphology and

molecules one has to have coded the characters right (ie delimited states

appropriately) and one has to use an appropriate model of evolution when fitting the

variation to the tree (see eg Omland 1999 Stevens 2006b) If the distributions of

apomorphies below are compared with those in other studies (eg Turgeon et al 2001

Bremer et al 2001 Endress 2001 Albach et al 2001a Judd amp Olmstead 2004 D Soltis

et al 2005b Zhang et al 2006 Judd et al 2007) the reader may find a number of

differences Although I have integrated such studies as far as is possible there are five

reasons that link to the preconditions just mentioned why there may be differences

where features are placed on trees

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

24

1 I may not have found all the information about a particular character there may be

disagreement over its interpretation or I have added information

2 The sampling of nearly all molecular studies is very incomplete (see Salisbury amp Kim

2001 for problems caused by sampling) indeed it is commonplace to decry the

incompleteness of molecular sampling But not only is the sampling in molecular studies

often less than we might wish that of the morphological and chemical characters whose

evolution we are interested in understanding is also often very poor So for many

anatomical chemical and embryological characters that are confidently said to

characterise families and other groups we all too often have no idea if those characters

are applicable to the whole clade or just to a subgroup within that clade However as

our sampling improves we can locate changes on the tree more precisely if in

unexpected positions Thus is is sometimes suggested that Ericaceae have ellagic acid

(Soltis et al 2005b) in fact the little evidence we have indicates that only a small clade

of some 80 species has ellagic acid and the rest of the family - itself alone about one

third of all Ericales - does not Similarly Ericaceae are noted for their

ectendomycorrhizal associations but Enkianthus sister to all other Ericaceae

apparently lacks these (Abe 2005) Thus these mycorrhizae and their associated

distinctive hair roots are unlikely to be apomorphies of Ericaceae - however it appears

that other families near Ericaceae may also have ectendomycorrhizae (see Asai 1934)

3 I am fitting characters to a very conservative tree with many polytomies although the

nodes that are utilised are for the most part strongly supported polytomies make the

optimisation of characters that is the assigment of character state change to a

particular node on the tree notably difficult (eg Madison amp Madison 2002) In nearly

all studies of the evolution of characters distributions of characters are optimised on a

more or less fully resolved tree and the construction of supertrees may yield yet more

detailed hypotheses of relationships (for literature on supertrees see Cotton amp

Wilkinson 2007 2008) Of course some nodes on such fully resolved trees andor

supertrees may have little support and optimisations of characters on such trees may

carry correspondingly little conviction Even parts of some of the trees used here have

poor support eg relationships within aquatic Alismatales etc although I indicate

when this is the case

4 Exactly how one goes about optimising a character on a tree is critically important

Even using simple parsimony optimisations (ACCTRAN or DELTRAN ACCelerated

TRANsitions or DELayed TRANsitions) the position of synapomorphies on trees - and

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

25

hence our ideas of evolution - may differ and this is still more true if one uses maximum

likelihood or Bayesian methods and makes apparently reasonable suggestions about

weighting gains over losses (eg Donoghue amp Ackerley 1996 Cunningham et al 1998

Omland 1997 1999 Ree amp Donoghue 1999 Polly 2001 Webster amp Purvis 2001

Ronquist 2004 Crisp amp Cook 2005) Sannier et al (2007) show how in in Arecaceae that

where on a phylogenetic tree one might peg changes in microsporogenesis will depend

on the methods one uses to do this and Pedersen et al (2007) discuss the sometimes

very substantial effect of node support on the posterior probabilities of ancestral

character states Here I use parsimony optimization not always as explicit as it might be

but I have often indicated where there are particularly important uncertainties as to the

positions of particular character changes on the tree

5 Finally although I have paid quite a lot of attention to the delimitation of the

character states that make up all the characterizations I have not spent enough time on

this critical operation If we are interested in understanding evolution then fitting the

basic variation - not character states - to a tree in principle allows greater flexibility in

understanding morphology in the context of local phylogenies (see also Stevens 2000

Endress 2005c) However many character states used here are delimited globally that

is they are circumscribed in the context of the variation shown by individual characters

across all angiosperms andor in the context of classic ideas of character evolution

Character states often have arbitrary limits and serve best to communicate

information whether they are in fact suitable for either phylogenetic analysis or

understanding evolution are separate issues Studies have rather unsurprisingly

perhaps but importantly shown that dividing the one character into different sets of

states may yield differing ideas of evolution of that character (eg Lamb Frye amp Kron

2003 Hibbett 2004) When looking at trees on which character states are optimised

one should bear in mind the problems surrounding the delimitation of states (eg

Stevens 2000 2006b) and the danger of using pollen or other types - constructs

based on many characters that vary independently but which effectively get lost in

these types thus Blackmore et al (2009) decomposed the pollen types in Asteraceae

into 52 characters

I use Remanes three main criteria of homology or better similarity when

determining the basic similarity of structures on different organisms (see Remane 1952)

These criteria are special properties position and intermediates Special properties

include anatomical or chemical characters gene expression data etc Position refers

to the position of an organ with respect to landmarks on the plant Although plants are

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

26

plants and landmarks are not as fixed as one might like trying to work out the relative

positions of parts is a good way of understanding morphology so long as one does not

try to out-do Procrustes Intermediates include intermediates found on the same

plant or on different plants Intermediates may observed during development or be

adult structures Thus one may find changes during development which suggest what

the nature of a particular structure is and very different stuctures on different plants

may turn out to be similar early in development Or one can simply compare different

taxa and see that structures that appear to be very different when comparing two

different taxa can be linked morphologically by looking at other taxa

Nevertheless the use of these criteria may not yield an unambiguous answer as to

what a structure is even given a solid phylogeny and an improved understanding of

development (see Jaramillo amp Kramer 2007 for a useful discussion) As Endress (2005c)

observed a number of features - position function development shape anatomy

histology gene activity and relationships to other taxa that clearly have petals - can be

used to distinguish a petal (for example) from other floral structures if a petal does not

have one of these features is it thereby not a petal Thus Maturen et al (2005) recently

found that floral organ diversity genes (B and C) were expressed in the large white

inflorescence bracts of Cornus (see also Costa et al 2005) Peney et al (2005) noted that

not all monosulcate pollen grains in monocots have the same developmental pathway

and that as a result such pollen might not have the same ancestral state Reeves and

Olmstead (2003) suggested that the genetic mechanisms causing monosymmetry in

Lamiales and Solanales were different and Serna and Martin (2006) described similar

problems with the development of hairs in Arabidopsis when compared with that of

hairs in Antirrhinum and Solanaceae Indeed as one perhaps might expect delimitation

of states and characters does not necessarily become easier with increasing knowledge

of development etc Thus Buzgo et al (2004) Matthews and Endress (2005) and others

have shown how hard it can be to distinguish between eg prophylls and other floral

structures as their behaviour is studied during the course of the development of the

flower Similarly at what concentration is a particular secondary metabolite deemed to

be present (Waterman (2007)

The validity of the approach used here that of fitting morphological variation to a

largely molecular-based tree may be questioned However I think it rather unlikely that

well-supported molecular branches will be overturned by morphological data Indeed

analyses of morphological data alone do provide support for many of the clades evident

in molecule-only analyses and analyses with morphological and molecular data

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

27

together may lead to increased support for clades (eg Hufford 1992 Nandi et al 1998

[but adding morphological data reduces support for a number of critical clades too]

Doyle amp Endress 2000 however in none of these papers is the use of morphology

without ambiguity) It is unfortunately clear that the use of morphology alone may not

suggest problems in the phylogenetic placement of taxa that later turn out to have been

wrongly included (eg Zhang et al 1992) Thus I have been wary of putting much weight

on clades that have only morphological support but note that I have treated molecule-

based clades with low bootstrap or jacknife support values (esp below 70) or low

posterior probabilies (below 095) likewise Although I may have been mistaken in

placing so much emphasis on molecular data in terms of providing the basic

phylogenetic framework for angiosperms morphological and molecular data are only

very rarely in irreconcilably strong conflict There are indeed a few places where the

conflict seems extreme These include the relative positions of the Monimiaceae and

Hernandiaceae (Laurales) the position of Hanguanaceae (Commelinales [as here] or

Zingiberales) and of Triplostegia (is it in Dipsacaceae or Valerianaceae - see

Dipsacales) Fossils are unlikely to affect the topologies of the trees presented here but

see below for their importance in understanding morphological evolution in general and

the evolution of angiosperms in particular Nevertheless some largely reject the idea

that trees based on molecular data alone can recover phylogenetic relationships

especially when branching points are old and prefer to used trees based on analysis of

morphological data including those taken from fossils (Hilton amp Bateman 2006 Farjon

2007)

All in all however the extent of the congruence between morphological and

molecular data is impressive and heartening and many clades can be characterised

morphologically It seemed in 1998 that there were no unambiguous morphological

synapomorphies for angiosperm orders (K Bremer 2000) and this is still true if by

unambiguous is meant non-homoplasious However many orders and other clades

have synapomorphies even if these may be indistinguishable at least at the current

level of morphological and developmental knowledge from parallel occurences

elsewhere As our morphological knowledge increases so too does the number of

apomorphies

Endress and Matthews (2006a) emphasize the importance of tendencies and

developmental constraints when thinking about characters of clades within the rosids -

many characters or character combinations occur in a rather sporadic fashion within

the clade and apparently notably less frequently outside it There are a number of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

28

examples of tendencies here and in the individual order pages such as the features

enclosed by parentheses in groups above the level of family eg see (cuticular waxes

as aggregated rodlets) for the commelinids and the discussion of the distribution of

polyandry that is flowers with numerous stamens in the asterid I + II groups etc

Tendencies may involve more than single characters When thinking of the

associations of features like integument thickness and vascularization of the integument

(Endress amp Matthews 2006a) one can imagine fairly simple developmental

preconditions being operative As Endress and Matthews (2006a) note it is difficult to

imagine vascular tissue developing in an integument only two cells thick Changes here

would be loosely correlated if morphologically linked However the strongly correlated

changes noted by Givnish et al (2005) are ecologically linked but are presumably

morphologicallydevelopmentally independent When there seem to be characters

evolving more or less together tests can be carried out to see if the changes are

concentrated on certain branches of the tree (eg Maddison 1990 Sanderson 1991

Maddison amp Maddison 2000) Such tests have rarely been carried out at higher levels in

angiosperms

One of the most striking examples of a tendency is the distribution of N-fixation

restricted as it is to a monophyletic group of four clades although it has arisen seven (or

perhaps several more) times independently within the N-fixing clade and several

members of two quite different kinds of bacteria are involved (eg Clawson et al 2004

Elliott et al 2007 Sprent amp James 2007) Other examples of work that bears on the issue

of tendencies include the findings that flowers of polysymmetrical Arabidopsis have

genes like TCP1 that are expressed asymmetrically during early development and TCP1

is a probable orthologue of the well-known CYC gene of Antirrhinum that is involved in

the development of monosymmetric slowers there (Cubas et al 2001 Costa et al 2005

etc) Parallelism might build on this underlying morphologically cryptic monosymmetry

even if details of the genetic mechanisms causing the monosymmetry evident in

particular groups may be different (Reeves amp Olmstead 2003 Cubas 2004 see above)

Similarly there have been several recent suggestions that the capability to synthesise a

particular metabolite may be switched off but not lost and so can sometimes be

reacquired (eg Wink amp Witte 1983 Wink 2003 Liscombe et al 2005 Larsson 2007

Waterman 2007) Hence perhaps the rather spotty distribution of many secondary

metabolites like ellagic acid the indole alkaloid camptothecin iridoids etc when

considered in the context of phylogenies As a non-botanical example - but a rather nice

one - Salwini-Plawen and Mayr (1961) suggested some time ago that there has been

considerable parallelism (40-65 or more independent origins) in the evolution of eyes in

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

29

metazoans However Pax 6 seems to be a master control gene that is involved in eye

formation perhaps throughout all bilateralians (eg Gehring amp Ikeo 1999 Erwin amp

Davidson 2002) which changes the nature of the problem Note however that in

general our invocation of evolutionary tendencies to explain the patterns of

distributions of characters is really an admission that we do not understand those

patterns

As we find out more about variation we will probably find fewer and fewer features

constant throughout a group Most unqualified statements of presence and absence

should properly be qualified as usually present or usually absent if one is thinking of

the characterisations on the order pages as encompassing the total variation within a

clade Thus Pistia alone among monocots as so far known has sieve tube plastids with

starch grains not protein crystals However this does not affect the fact that sieve tube

plastids with cuneate protein crystals and lacking starch grains are an apomorphy for

monocots Furthermore fossils in a number of cases suggest character combinations

unknown in extant taxa as may be seen in the discussions of Fagaceae Platanaceae

Iteaceae Calycanthaceae etc Confusing the issue there may be questions as to where

exactly on the tree a particular fossil is to be placed (eg see Nymphaeaceae

Calycanthaceae Archaefructus etc)

To summarize given our current understandings of both phylogenies and

characters evolution of some characters in which we are interested seems very labile

(see eg D Soltis et al 2005b Endress and Matthews 2006a Stevens 2006b Ekman et

al 2008 for a good recent study) and I have been cautious when talking about character

evolution Much effort must continue to be spent in summarizing characters of well-

established clades at all levels providing features by which they may be recognized and

signaling synapomorphies Remember that (1) the basic morphological anatomical and

chemical knowledge of many critical taxa is woefully incomplete (2) different

assumptions about character evolution may greatly affect the position of

synapomorphies on trees (3) in many cases relationships within and between many

groups are too uncertain at present to worry very much about synapomorphies and

(4) we must be clear about what we do and do not not know As mentioned above I

have indicated in a number of places where there are particularly important

uncertainties as to where characters should be placed on the tree Nevertheless it is a

relatively easy matter to update notes such as these and it can be a simple matter to

incorporate new data on characters that have never before been considered in the

context of a tree Much basic - and unfortunately perhaps unfashionable - work must be

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

30

carried to clarify the distribution of morphological anatomical and chemical characters

for examples of what can to be done see the work of P K Endress and collaborators

(floral morphology and development) and S R Jensen and collaborators (iridoids) But

acquiring information about nodal anatomy cell and tissue distribution nodal anatomy

and the like is not difficult at all and should be generally encouraged

But all this is merely a necesary prelude to the understanding of evolution For this

one needs to know a lot more including dating the phylogeny understanding the

function(s) of characters factoring in relevant aspects of the palaeoenvironment in

which an apomorphy first appeared etc

SUMMARY OF THE SYSTEM

Below is a formalised summary of the relationships within orders of the families of seed

plants There are a few families that are not recognised even as options in APG II and

vice versa as well as a few extra orders The families to be recognised in the most

recent edition of Mabberleys The Plant Book (Mabberley 2008) are also largely

consistent with those below But all differences are trivial and will - I hope - eventually

disappear

Square brackets - [] - enclose clades the plus sign - + - designates sister taxa a

comma - - denotes part of a polytomy and quotation marks - - denotes a

paraphyletic group

SEED PLANTS

GYMNOSPERMS

Cycadales

Cycadaceae + Zamiaceae

Ginkgoales

Ginkgoaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

31

Pinales

Pinaceae [[Araucariaceae [Podocarpaceae + Phyllocladaceae]] [Sciadopityaceae

[Taxaceae + Cupressaceae]]]

Gnetales

Ephedraceae [Gnetaceae + Welwitschiaceae]

ANGIOSPERMSFLOWERING PLANTS

Amborellales

Amborellaceae

Nymphaeales

Hydatellaceae [Cabombaceae + Nymphaeaceae]

Austrobaileyales

Austrobaileyaceae [Schisandraceae + Trimeniaceae]

Chloranthales

Chloranthaceae

MAGNOLIIDS

Magnoliales

Myristicaceae [Magnoliaceae [[Himantandraceae + Degeneriaceae] [Eupomatiaceae +

Annonaceae]]]

Laurales

Calycanthaceae [[Siparunaceae [Gomortegaceae + Atherospermataceae]] [Monimiaceae

[Hernandiaceae + Lauraceae]]]

Canellales

Canellaceae + Winteraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

32

Piperales

[Hydnoraceae + Aristolochiaceae] [Piperaceae + Saururaceae]

MONOCOTS

Acorales

Acoraceae

Alismatales

Araceae [Tofieldiaceae [[Alismataceae [Hydrocharitaceae + Butomaceae]]

[Scheuchzeriaceae [Aponogetonaceae [Juncaginaceae [Maundiaceae [[Posidoniaceae

[Ruppiaceae + Cymodoceaceae]] [Zosteraceae + Potamogetonaceae]]]]]]]

Petrosaviales

Petrosaviaceae

Dioscoreales

Nartheciaceae [[Taccaceae + Thismiaceae] [Burmanniaceae + Dioscoreaceae]]

Pandanales

Velloziaceae Triuridaceae Stemonaceae [Pandanaceae + Cyclanthaceae]

Liliales

Corsiaceae [Campynemataceae [Petermanniaceae [Colchicaceae + Alstroemeriaceae]]

Melanthiaceae [[Philesiaceae + Rhipogonaceae] [Smilacaceae + Liliaceae]]]

Asparagales

Orchidaceae [[Boryaceae [Blandfordiaceae [Lanariaceae [Asteliaceae + Hypoxidaceae]]]]

[[Ixioliriaceae + Tecophilaeaceae] [Doryanthaceae [Iridaceae [Xeronemataceae

[Xanthorrhoeaceae [Alliaceae + Asparagaceae]]]]]]]

COMMELINIDS

Unplaced

Dasypogonaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

33

Arecales

Arecaceae

Poales

[Typhaceae + Bromeliaceae] [Rapateaceae [[Thurniaceae [Juncaceae + Cyperaceae]]

[[Mayacaceae [Eriocaulaceae + Xyridaceae]] [Flagellariaceae [Anarthriaceae

[Centrolepidaceae + Restionaceae]]] [Joinvilleaceae [Ecdeiocoleaceae + Poaceae]]]]]]]

Commelinales

[Commelinaceae + Hanguanaceae] [Philydraceae [Haemodoraceae + Pontederiaceae]]

Zingiberales

Musaceae [Strelitziaceae + Lowiaceae] Heliconiaceae [[Cannaceae + Marantaceae]

[Costaceae + Zingiberaceae]]

Ceratophyllales

Ceratophyllaceae

EUDICOTS

Ranunculales

Eupteleaceae [Papaveraceae [[[Lardizabalaceae + Circaeasteraceae] [Menispermaceae

[Berberidaceae + Ranunculaceae]]]]

Sabiales

Sabiaceae

Proteales

Nelumbonaceae [Platanaceae + Proteaceae]

Trochodendrales

Trochodendraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

34

Buxales

Haptanthaceae [Buxaceae + Didymelaceae]

CORE EUDICOTS

Gunnerales

Gunneraceae + Myrothamnaceae

Dilleniales

Dilleniaceae

Saxifragales

Peridiscaceae [[Paeoniaceae [Altingiaceae [Hamamelidaceae [Cercidiphyllaceae +

Daphniphyllaceae]]]] [[Crassulaceae [Aphanopetalaceae [Tetracarpaeaceae

[Penthoraceae + Haloragaceae]]]] [Iteaceae [Grossulariaceae + Saxifragaceae]]]]

Cynomoriaceae unplaced

Vitales

Vitaceae

ROSIDS

FABIDROSID I

Zygophyllales

Krameriaceae + Zygophyllaceae

Celastrales

Lepidobotryaceae + Celastraceae

Oxalidales

Huaceae [[Connaraceae + Oxalidaceae] [Cunoniaceae [Elaeocarpaceae [Brunelliaceae +

Cephalotaceae]]]]

Malpighiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

35

[Achariaceae [Goupiaceae [Violaceae + Passifloraceae] [Lacistemataceae Salicaceae]]

[Lophopyxidaceae Putranjivaceae] [Ctenolophonaceae [Erythroxylaceae +

Rhizophoraceae]] Linaceae Ixonanthaceae Humiriaceae Irvingiaceae

Centroplacaceae Pandaceae [[Bonnetiaceae + Clusiaceae] [Calophyllaceae

[Hypericaceae + Podostemaceae]]] [Malpighiaceae + Elatinaceae] Ochnaceae

[Peraceae [Rafflesiaceae + Euphorbiaceae]] [Phyllanthaceae + Picrodendraceae]

[Balanopaceae [[Trigoniaceae + Dichapetalaceae] [Chrysobalanaceae +

Euphroniaceae]]] Caryocaraceae

N-FIXING CLADE

Fabales

Quillajaceae [Fabaceae [Polygalaceae + Surianaceae]]

Rosales

Rosaceae [[Barbeyaceae [Elaeagnaceae Dirachmaceae Rhamnaceae]] [Ulmaceae

[Cannabaceae [Moraceae + Urticaceae]]]]

Cucurbitales

Anisophylleaceae [[Corynocarpaceae + Coriariaceae] [Cucurbitaceae [Tetramelaceae

[Datiscaceae + Begoniaceae]]]] Apodanthaceae

Fagales

Nothofagaceae [Fagaceae [[Myricaceae + Juglandaceae] [Casuarinaceae

[Ticodendraceae + Betulaceae]]]]

MALVIDROSID II

pgtGeraniales

Geraniaceae [[Melianthaceae + Francoaceae] [Vivianaceae + Ledocarpaceae]]

Myrtales

Combretaceae [[Onagraceae + Lythraceae] [[Vochysiaceae + Myrtaceae]

[Melastomataceae [Crypteroniaceae [Alzateaceae + Penaeaceae]]]]]

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

36

Crossosomatales

[Staphyleaceae [Guamatelaceae [Crossosomataceae + Stachyuraceae]]] [Aphloiaceae

[Geissolomataceae + Strasburgeriaceae]]

Picramniales

Picramniaceae

Sapindales

Biebersteiniaceae Nitrariaceae [Kirkiaceae [Anacardiaceae + Burseraceae]]

[Sapindaceae [Simaroubaceae Rutaceae Meliaceae]]

Huerteales

Gerradinaceae [Dipentodontaceae + Tapisciaceae]

Malvales

Neuradaceae [Thymelaeaceae [Sphaerosepalaceae Bixaceae [Cistaceae

[Sarcolaenaceae + Dipterocarpaceae]] [Cytinaceae + Muntingiaceae] Malvaceae]]

Brassicales

[Akaniaceae + Tropaeolaceae] [[Moringaceae + Caricaceae] [Setchellanthaceae

[Limnanthaceae [[Koeberliniaceae [Bataceae + Salvadoraceae]] [Emblingiaceae

[Pentadiplandraceae [Gyrostemonaceae Resedaceae Stixis and relatives] Tovariaceae

[Capparaceae [Cleomaceae + Brassicaceae]]]]]]]]

Berberidopsidales

Aextoxicaceae + Berberidopsidaceae

Santalales

Erythropalaceae Olacaceae [[Loranthaceae [Misodendraceae + Schoepfiaceae]]

[Opiliaceae + Santalaceae]] Balanophoraceae unplaced

Caryophyllales

[[Droseraceae [Nepenthaceae [Drosophyllaceae [Ancistrocladaceae +

Dioncophylleaceae]]]]] [[Frankeniaceae + Tamaricaceae] [Polygonaceae +

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

37

Plumbaginaceae]]] [Rhabdodendraceae [Simmondsiaceae [[Asteropeiaceae +

Physenaceae] [[Caryophyllaceae [Achatocarpaceae + Amaranthaceae]]

[Stegnospermataceae [Limeaceae [[Lophocarpaceae [Barbeuiaceae [Aizoaceae

[Sarcobataceae Phytolaccaceae Nyctaginaceae]]]] [Molluginaceae [Halophytaceae

Montiaceae Basellaceae Didiereaceae [Talinaceae [Portulacaceae [Anacampseros etc +

Cactaceae]]]]]]]]]]]]]

ASTERIDS

Cornales

[Cornaceae + Nyssaceae] [Hydrangeaceae + Loasaceae] Hydrostachyaceae [Curtisiaceae

+ Grubbiaceae]

Ericales

[Balsaminaceae [Marcgraviaceae + Tetrameristaceae]] [[Polemoniaceae +

Fouquieraceae] Lecythidaceae [[Sladeniaceae + Pentaphylacaceae] [Sapotaceae

[Ebenaceae [Maesaceae [Theophrastaceae [Myrsinaceae + Primulaceae]]]]]

[Mitrastemonaceae Theaceae [Symplocaceae [Styracaceae + Diapensiaceae]]

[[Sarraceniaceae [Roridulaceae + Actinidiaceae] [Clethraceae [Cyrillaceae +

Ericaceae]]]]]]

LAMIIDASTERID I

Unplaced

Metteniusaceae Oncothecaceae Icacinaceae and some unplaced ex-Icacinaceae

Garryales

Garryaceae + Eucommiaceae

Unplaced

Boraginaceae Vahliaceae

Gentianales

Rubiaceae [Gentianaceae [Loganiaceae [Gelsemiaceae + Apocynaceae]]]

Lamiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

38

Plocospermataceae [Oleaceae [Tetrachondraceae [Calceolariaceae

[Gesneriaceae Peltanthera [Pedaliaceae Carlemanniaceae Martyniaceae Stilbaceae

Plantaginaceae Linderniaceae Bignoniaceae Verbenaceae Lamiaceae Paulowniaceae

Schlegeliaceae Thomandersiaceae Phrymaceae Rehmannia Acanthaceae

Scrophulariaceae Orobanchaceae Byblidaceae Lentibulariaceae]]]]]

Solanales

[Montiniaceae [Sphenocleaceae + Hydroleaceae]] [Convolvulaceae + Solanaceae]

CAMPANULIDASTERID II

Aquifoliales

[[Cardiopteridaceae + Stemonuraceae] [Aquifoliaceae [Helwingiaceae +

Phyllonomaceae]]

Asterales

[Rousseaceae + Campanulaceae] Pentaphragmataceae [[Alseuosmiaceae [Phellinaceae

+ Argophyllaceae]] Stylidiaceae [Menyanthaceae [Goodeniaceae [Calyceraceae +

Asteraceae]]]]

Escalloniales

Escalloniaceae

Bruniales

[Bruniaceae + Columelliaceae]

Apiales

Pennantiaceae [Torricelliaceae [Griseliniaceae [Pittosporaceae [Araliaceae

[Myodocarpaceae + Apiaceae]]]]]

Paracryphiales

Paracryphiaceae

Dipsacales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

39

Adoxaceae [Diervillaceae [Caprifoliaceae [Linnaeaceae [Morinaceae [Dipsacaceae +

Valerianaceae]]]]]

LINKS TO ORDERS AND FAMILIES

[Back to Top]

MAIN GROUPINGS

Asterids commelinids core eudicots asterid 1 asterid

2 eudicots gymnosperms Magnoliophyta monocots N-fixing claderosids seed plants

ALPHABETICAL LISTING OF ALL ORDINAL NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | L | M | N | O | P | Q | R | S | T | U | V | W | X | Z

Abietales Acanthales Acerales Acorales Actinidiales Actinostrobales Adoxales Aescul

ales Agavales Aizoales AkanialesAlismatales Alliales Alseuosmiales Alstroemeriales

Altingiales Amaranthales Amaryllidales Amborellales AmbrosialesAmmiales Amomal

es Ancistrocladales Anisophylleales Annonales Anthobolales Apiales Apocynales Apo

nogetonalesAquifoliales Arales Araliales Aralidiales Araucariales Arecales Aristolochi

ales Athrotaxidales Asarales AsclepiadalesAsparagales Asphodelales Asparagales Ast

eliales Atriplicales Aucubales Austrobaileyales Avenales

Balanitales Balanopales Balanophorales Balsaminales Barbeyales Barclayales Batales

Begoniales BerberidalesBerberidopsidales Betulales Biebersteiniales Bignoniales Bix

ales Boraginales Brassicales Brexiales Bromeliales BrunialesBrunoniales Burmannial

es Burserales Butomales Buxales Byblidales

Cactales Callitrichales Calycanthales Calycerales Campanulales Campynematales Can

ellales Cannales CapparalesCaprifoliales Cardiopteridales Carduales Caricales Carle

manniales Caryophyllales Cassiales Casuarinales CelastralesCentrolepidales Cephalot

ales Cephalotaxales Ceratophyllales Cercidiphyllales Chenopodiales Chironiales Chlor

anthalesChrysobalanales Cinchonales Circaeasterales Cistales Citrales Cocosales Col

chicales Columelliales CombretalesCommelinales Connarales Convolvulales Coriarial

es Cornales Corylales Corynocarpales Crassulales CrossosomatalesCucurbitales Cunn

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

40

inghamiales Cunoniales Cupressales Cycadales Cyclanthales Cymodoceales Cynarales

CynomorialesCyperales Cyrillales Cytinales

Daphnales Daphniphyllales Dasypogonales Datiscales Degeneriales Desfontainiales D

iapensiales Didymelales DillenialesDioales Dioncophyllales Dioscoreales Diospyrales

Dipentodontales Dipsacales Droserales

Ebenales Echiales Elaeagnales Elaeocarpales Elatinales Elodeales Emmotales Empetr

ales Ephedrales Ericales EriocaulalesErythropalaless Escalloniales Eucommiales Euph

orbiales Eupomatiales Eupteleales Euryalaless

Fabales Fagales Falcatifoliales Ficales Flacourtiales Flagellariales Fouquieriales Franc

oales Frangulales

Galiales Garryales Geissolomatales Gentianales Geraniales Gesneriales Ginkgoales G

laucidiales Globulariales GnetalesGoodeniales Greyiales Griseliniales Grossulariales

Grubbiales Gunnerales Gyrocarpales Gyrostemonales

Haemodorales Haloragales Hamamelidales Hanguanales Heisteriales Helleborales He

lwingiales Himantandrales HippuridalesHomaliales Hortensiales Huales Huerteales

Hydatellales Hydnorales Hydrangeales Hydrastidales HydrocharitalesHydropeltidales

Hydrostachyales Hypericales Hypoxidales

Icacinales Illiciales Iridales Irvingales Iteales Ixiales

Jasminales Juglandales Julianiales Juncaginales Juncales

Lacistematales Lactoridales Lamiales Lardizabalales Laurales Lecythidales Ledocarpal

es Leitneriales LentibularialesLigustrales Liliales Limnanthales Linales Liriales Loasal

es Lobeliales Loganiales Lonicerales Loranthales LowialesLythrales

Magnoliales Malpighiales Malvales Marathrales Marcgraviales Mayacales Medusagy

nales Medusandrales MelanthialesMelastomatales Meliales Melianthales Meliosmal

es Menispermales Menyanthales Metteniusales MitrastemonalesMiyoshiales Moni

miales Moringales Musales Myricales Myristicales Myrothamnales Myrsinales Myrta

les

Najadales Nandinales Narcissales Nartheciales Nelumbonales Nepenthales Neuradal

es Nitrariales Nolanales NothofagalesNyctaginales Nymphaeales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

41

Ochnales Oenotherales Olacales Oleales Onagrales Oncothecales Opuntiales Orchid

ales Oxalidales

Saxifragales Pandales Pandanales Papaverales Paracryphiales Parasitaxales Paridales

Parnassiales PassifloralesPenaeales Pennantiales Pentaphragmatales Peridiscales P

etiveriales Petrosaviales Phellinales Philydrales PhyllanthalesPhysenales Phytolaccal

es Picramniales Pinales Pinguiculales Piperales Pittosporales Plantaginales Platanale

sPlumbaginales Poales Podocarpales Podophyllales Podostemales Polemoniales Pol

ygalales Polygonales PontederialesPortulacales Posidoniales Potamogetonales Primu

lales Proteales

Quercales Quillajales Quintiniales

Rafflesiales Ranunculales Rapateales Resedales Restionales Rhabdodendrales Rhamn

ales Rhinanthales RhizophoralesRhodorales Rhoipteleales Roridulales Rosales Rouss

eales Rubiales Ruppiales Rutales

Sabiales Salicales Salvadorales Samolales Samydales Sanguisorbales Santalales Sapin

dales Sapotales SarracenialesSaxegotheales Saxifragales Scheuchzeriales Sciadopitya

les Scleranthales Scrophulariales Scyphostegiales SedalesSilenales Simmondsiales S

milacales Solanales Sphenocleales Sphenostemonales Stangeriales Stellariales Stemo

nalesStilbales Stylidiales Styracales Surianales

Taccales Tamales Tamaricales Taxales Taxodiales Tecophilaeales Terebinthales Tern

stroemiales Theales TheligonalesThymelaeales Tiliales Tofieldiales Torricelliales Tov

ariales Tribelales Trilliales Trimeniales Triuridales TrochodendralesTropaeolales Turn

erales Typhales

Ulmales Urticales

Vacciniales Vahliales Vallisneriales Velloziales Veratrales Verbenales Viburnales Vinc

ales Violales Viscales VitalesVochysiales

Welwitschiales Winterales

Xanthorrhoeales Ximeniales Xyridales

Zamiales Zingiberales Zosterales Zygophyllales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

42

ALPHABETICAL LISTING OF ALL FAMILY NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | K | L | M | N | O | P | Q | R | S | T | U | V | W | X | Y

|Z

Back to Home

In some cases clicking on a family synonym will send you directly to the subfamily in

which the genus of the basionym is to be found

Abietaceae Abolbodaceae Abrophyllaceae Acalyphaceae Acanthaceae [Acanthochla

mydaceae] Acarnaceaee AceraceaeAchariaceae Achatocarpaceae Achradaceae Achy

ranthaceae Acmopylaceae Acoraceae Acristaceae ActaeaceaeActinidiaceae Actinota

ceae Amentotaxaceae Adenogrammaceae Adoxaceae Aegialitidaceae Aegicerataceae

Aegilopaceae Aeginetiaceaee Aegiphilaceae Aesculaceae Aextoxicaceae Agapanthac

eae Agavaceae AgdestidaceaeAgialidaceae Agrimoniaceae Agrostidaceae Ailanthace

ae Aitoniaceae Aizoaceae Akaniaceae Alangiaceae AlchemillaceaeAldrovandaceae A

lismataceae Alliaceae Allioniaceae Allophylaceae Aloaceae Alopecuraceae Alpiniacea

e AlseuosmiaceaeAlsinaceae Alsodeiaceae Alstroemeriaceae Altingiaceae Alzateacea

e Amaranthaceae Amaryllidaceae Amborellaceae Ambrosiaceae Amentotaxaceae A

mmanniaceae Ammiaceae Amomaceae Ampelidopsaceae Amygdalaceae Amyridacea

eAnacardiaceae Anagallidaceae Anarthriaceae Anchusaceae Ancistrocladaceae Andr

omedaceae AndropogonaceaeAndrostachyaceae Androsynaceae Anemarrhenaceae

Anemonaceae Angelicaceae Anisophylleaceae AnnonaceaeAnomochloaceae Anopter

aceae Anrederaceae Anthemidaceae Anthericaceae Anthobolaceae Antidesmataceae

Antirrhinaceae Antoniaceae Aparinaceae Aphanopetalaceae Aphloiaceae Aphyllanth

aceae Apiaceae ApocynaceaeApodanthaceae Aponogetonaceae Aporusaceae Apose

ridaceae Apostasiaceae Aptandraceae AquifoliaceaeAquilariaceae Aquilegiaceae A

raceae Aragoaceae Araliaceae Aralidiaceaee Araucariaceae Arbutaceae Arceuthidace

aeArceuthobiaceae Arctostaphylaceae Arctotidaceae Ardisiaceae Arecaceae Argoph

yllaceae Arisaraceae AristolochiaceaeAristoteliaceae Arjonaceae Armeriaceae Artem

isiaceae Arthrotaxidaceae Artocarpaceae Arundinaceae ArundinellaceaeAsaraceae A

sclepiadaceae Ascyraceaee Asiraceae Asparagaceae Asperulaceae Asphodelaceae As

pidistraceaeeAsteliaceae Asteraceae Asteranthaceae Asteropeiaceae Astragalaceae

Astrocarpaceae AthanasiaceaeAtherospermataceae Atriplicaceae Atropaceae Aucuba

ceae Aurantiaceae Austrobaileyaceae Austrotaxaceae AvenaceaeAverrhoaceae Avet

raceae Avicenniaceae Azaleaceae Azimaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

43

Balanitaceae Balanopaceae Balanophoraceae Balsameaceae Balsaminaceae Bambusa

ceae Barbaceniaceae BarbeuiaceaeBarbeyaceae Barclayaceae Barringtoniaceae Base

llaceae Bataceae Baueraceae Bauhiniaceae Baxteriaceae BegoniaceaeBehniaceae B

elangeraceae Belloniaceae Bembiciaceae Berberidaceae Berberidopsidaceae Berryac

eaee BersamaceaeBertyaceae Berzeliaceae Besleriaceae Betaceae Betulaceae Biebe

rsteiniaceae Bifariaceae Bignoniaceae BischofiaceaeBixaceae Blakeaceae Blandfordia

ceae Blattiaceae Blepharocaryaceae Blitaceae Blyxaceae Boerlagellaceae Bolivaracea

eBombacaceae Bonnetiaceae Bontiaceae Boopidaceae Boraginaceae Borassaceae B

oroniaceae BoryaceaeBotryodendraceae Bougainvilleaceae Boweniaceae Brassicacea

e Brachycaulaceae Bretschneideraceae BrexiaceaeBromeliaceae Brownlowiaceae Br

unelliaceae Bruniaceae Brunoniaceae Brunsvigiaceae Bryoniaceae BuchneraceaeBuci

daceae Buddlejaceae Buglossaceae Bulbocodiaceae Bumeliaceae Bupleuraceae Burc

hardiaceae BurmanniaceaeBurseraceae Butneriaceae Butomaceae Buxaceae Byblida

ceae Byttneriaceae

Cabombaceae Cacaoaceae Cactaceae Caesalpiniaceae Caladiaceae Calamaceae Calce

olariaceae CalectasiaceaeCalendulaceae Callaceae Callicomaceae Calligonaceae Calli

traceae Callitrichaceae Calochortaceae CalophyllaceaeCalthaceae Calycanthaceae Ca

lyceraceae Cambogiaceae Camelliaceae Campanulaceae CampynemataceaeCanacom

yricaceae Canellaceae Cannabaceae Cannaceae Canopodaceae Canotiaceae Cansjera

ceae CapparaceaeCaprariaceae Caprifoliaceae Cardiopteridaceae Carduaceae Carica

ceae Carissaceae Carlemanniaceae CarpinaceaeCarpodetaceae Cartonemataceae Ca

ryocaraceae Caryophyllaceae Casearieaceae Cassiaceae CassipoureaceaeCassythacea

e Castaneaceae Castelaceae Casuarinaceae Catesbaeaceae Cathedraceae Cecropiace

ae CedraceaeCedrelaceae Celastraceae Celosiaceae Celtidaceae Centaureaceae Cen

trolepidaceae Centroplacaceae CepaceaeCephalanthaceae Cephalotaceae Cephalota

xaceae Cerastiaceae Ceratoniaceae Ceratophyllaceae CerberaceaeCercidiphyllaceae

Cercocarpaceae Cercodiaceae Cereaceae Cerinthaceae Ceroxylaceae Cestraceae Cev

alliaceaeChailletiaceae Chamaedoraceae Chamaemoraceae Chamelauciaceae Chaun

ochitonaceae Chelidoniaceae ChelonaceaeChenopodiaceae Chimonanthaceae Chingi

thamnaceae Chionographidaceae Chiranthodendraceae ChironiaceaeChlaenaceae Ch

loanthaceae Chloranthaceae Chloridaceae Chlorogalaceae Chrysobalanaceae Cicerac

eae CichoriaceaeCimicifugaceae Cinchoniaceae Circaeaceae Circaeasteraceae Cissac

eae Cistaceae Citraceae Clematidaceae CleomaceaeClethraceae Clusiaceae Cneorac

eae Cnestidaceae Cnicaceae Cobaeaceae Cochlospermaceae Cocosaceae Coffeaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

44

Colchicaceae Coleogynaceae Columelliaceae Combretaceae Commelinaceae Comocl

adiaceae Compositae CompsoaceaeConantheraceae Connaraceae Conostylidaceae C

onvallariaceae Convolvulaceae Cordiaceae CoreopsidaceaeCoriandraceae Coriariacea

e Coridaceae Corispermaceae Cornaceae Corokiaceae Coronillaceae Correaceae Cor

rigiolaceaeCorsiaceae Corydalaceae Corylaceae Corynocarpaceae Coryphaceae Cost

aceae Cotyledonaceae CoulaceaeCoutariaceae Coutoubiaceae Crassulaceae Crescen

tiaceae Cressaceae Crinaceae Crocaceae CroomiaceaeCrossosomataceae Crotonacea

e Cruciferae Cryptaceae Crypteroniaceae Cryptocorynaceae CryptomeriaceaeCtenol

ophonaceae Cucurbitaceae Cunninghamiaceae Cunoniaceae Cupressaceae Curcumac

eae Curtisiaceae CuscutaceaeCyananthaceae Cyanastraceae Cynanchaceae Cyanella

ceae Cycadaceae Cyclanthaceae CyclantheraceaeCyclocheilaceae Cydoniaceae Cymo

doceaceae Cynaraceae Cynocrambaceae Cynomoriaceae Cyperaceae CyphiaceaeCyp

hocarpaceae Cypripediaceae Cyrillaceae Cyrtandraceae Cyrtanthaceae Cytinaceae

Dacrycarpaceae Dactylanthaceae Damasoniaceae Daphnaceae Daphniphyllaceae Das

ypogonaceae DatiscaceaeDaturaceae Daucaceae Davidiaceae Davidsoniaceae Decais

neaceae Deeringiaceae Degeneriaceae DelphiniaceaeDendrophthoaceae Columelliac

eae Detariaceae Dialypetalanthaceae Dianellaceae Dianthaceae DiapensiaceaeDicha

petalaceae Dichondraceae Diclidantheraceae Dicrastylidiaceae Dictamnaceae Didiere

aceae DidymelaceaeDidymocarpaceae Diegodendraceae Diervillaceae Digitalidaceae

e Dilatridaceae Dilleniaceae Dioaceae DionaeaceaeDioncophyllaceae Dioscoreaceae

Diosmaceae Diospyraceae Dipentodontaceae Diphylleiaceae Diplolaenaceae Dipsaca

ceaeDipterocarpaceae Dirachmaceae Disanthaceae Diselmaceae Dodonaeaceae Do

mbeyaceae Donatiaceae DorsteniaceaeDortmannaceae Doryanthaceae Drabaceae D

racaenaceae Dracontiaceae Drimyidaceae Droseraceae DrosophyllaceaeDryadaceae

Duabangaceae Duckeodendraceae Dulongiaceae Durantaceae Durionaceae Dysphani

aceae

Ebenaceae Eccremidaceae Ecdeiocoleaceae Echiaceae Echinopaceae Ehretiaceae Ela

eagnaceae ElaeocarpaceaeElegiaceae Elatinaceae Ellisiaceae Ellisiophyllaceae Elodea

ceae Elytranthaceae Embeliaceae Emblingiaceae EmmotaceaeEmpetraceae Encepha

lartaceae Engelhardtiaceae Enhalaceae Epacridaceae Ephedraceae Ephemeranthacea

eEpilobiaceae Epimediaceae Eragrostidaceae Eremolepidaceae Eremosynaceae Erica

ceae Erinaceae EriocaulaceaeEriogonaceae Eriospermaceae Erodiaceae Erycibaceae

Eryngiaceae Erysimaceae Erythroniaceae ErythropalaceaeErythrospermaceae Erythro

xylaceae Escalloniaceae Eschscholziaceae Eucomidaceae Eucommiaceae Eucryphiace

aeEuonymaceae Eupatoriaceae Euphorbiaceae Euphrasiaceae Euphroniaceae Eupo

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

45

matiaceae Eupteleaceae EuryalaceaeEustrephaceae Euthemidaceae Eutocaceae Exb

ucklandiaceae Exocarpaceae

Fabaceae Fagaceae Falcatifoliaceae Ferulaceae Festucaceae Ficaceae Ficoidaceae Fit

zroyaceae FlacourtiaceaeFlagellariaceae Flindersiaceae Foetidiaceae Forestieraceae

Fothergillaceae Fouquieriaceae Fragariaceae FrancoaceaeFrangulaceae Frankeniacea

e Fraxinaceae Fraxinellaceae Freycinetiaceae Fritillariaceae Fuchsiaceae Fumariacea

eFunkiaceae

Gaiadendraceae Galacaceae Galanthaceae Galaxiaceae Galedupaceae Galeniaceae G

aliaceae Garciniaceae GardeniaceaeGardneriaceae Garryaceae Geissolomataceae Ge

itonoplesiaceae Gelsemiaceae Geniostomaceae GentianaceaeGeonomataceae Geosir

idaceae Geraniaceae Gerrardinaceae Gesneriaceae Gethyllidaceae Gilliesiaceae Gina

lloaceaeGinkgoaceae Gisekiaceae Gladiolaceae Glaucidiaceae Glechomaceae Glinace

ae Globulariaceae Gnaphaliaceae GnetaceaeGoetzeaceae Gomortegaceae Gomphiac

eae Gomphrenaceae Gonystylaceae Goodeniaceae Gordoniaceae GouaniaceaeGoup

iaceae Gramineae Gratiolaceae Grewiaceae Greyiaceae Grielaceae Griseliniaceae Gr

onoviaceae GrossulariaceaeGrubbiaceae Guaiacanaceae Guamatelaceae Guettardac

eae Gunneraceae Gustaviaceae Guttiferae GyrocarpaceaeGyrostemonaceae

Hachetteaceae Haemanthaceae Haemodoraceae Halesiaceae Hallieraceae Halocarpa

ceae Halophilaceae HalophytaceaeHaloragaceae Hamamelidaceae Hanguanaceae H

aptanthaceae Harmandaceae Hebenstretiaceae HectorellaceaeHederaceae Hedyosm

aceae Hedyotidaceae Hedysaraceae Heisteriaceae Heleniaceae Heliamphoraceae H

elianthaceaeHelianthemaceae Helichrysaceae Heliconiaceae Helicteraceae Heliotropi

aceae Helleboraceae Heloniadaceae HelosidaceaeHelwingiaceae Hemerocallidaceae

Hemimeridaceae Hemiphylacaceae Henriqueziaceae Henslowiaceae Hermanniaceae

Hernandiaceae Herniariaceae Herreriaceae Hesperocallidaceae Heterantheraceae He

teropyxidaceae HeterostylaceaeHibbertiaceae Hibiscaceae Hilleriaceae Himantandra

ceae Hippocastanaceae Hippocrateaceae HippomanaceaeHippophaeaceae Hippurida

ceae Hirtellaceae Holacanthaceae Homaliaceae Hopkinsiaceae Hoplestigmataceae H

ordeaceaeHornschuchiaceae Hortensiaceae Hortoniaceae Hostaceae Houstoniaceae

Huaceae Huerteaceae HugoniaceaeHumbertiaceae Humiriaceae Hyacinthaceae Hyd

atellaceae Hydnoraceae Hydrangeaceae Hydrastidaceae HydrillaceaeHydrocharitacea

e Hydrocotylaceae Hydrogetonaceae Hydroleaceae Hydropeltidaceae Hydrophylacac

eae HydrophyllaceaeHydrostachyaceae Hymenocardiaceae Hypecoaceae Hyperanth

eraceae Hypericaceae Hypopityaceae HypoxidaceaeHypseocharitaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

46

Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta

ceae ImperatoriaceaeInocarpaceae Inulaceae Ionidiaceae Iriarteaceae Iridaceae Irvi

ngiaceae Isnardiaceae Isophysidaceae IteaceaeStrasburgeriaceae Ixiaceae Ixioliriace

ae Ixonanthaceae

Jalapaceae Jamboliferaceae Japonoliriaceae Jasionaceae Jasminiaceae Johnsoniaceae

Joinvilleaceae JuglandaceaeJulianiaceae Juncaceae Juncaginaceae Juniperaceae Jus

siaeaceae Justiciaceae

Kadsuraceae Kaliphoraceae Kalmiaceae Kaniaceae Kiggelariaceae Kingdoniaceae Kin

giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae

Krameriaceae

Labiatae Lacandoniaceae Lachenaliaceae Lacistemataceae Lactoridaceae Lactucaceae

Lagerstroemiaceaee LamiaceaeLampsanaceae Lanariaceae Langsdorffiaceae Lantan

aceae Lapageriaceae Lardizabalaceae Lasiopetalaceae LauraceaeLawsoniaceae Laxm

anniaceae Lecythidaceae Ledaceae Ledocarpaceae Leeaceae Leguminosae Leitneriac

eae LemnaceaeLennoaceae Lentibulariaceae Lentiscaceae Leoniaceae Leonticaceae

Lepidobotryaceae LepidocarpaceaeLepidocaryaceae Lepidocerataceae Leptaulaceae

Leptospermaceae Lepturaceae Lepuropetalaceae LeuctenbergiaceaeLibocedraceae

Licaniaceae Ligustraceae Lilaeaceae Liliaceae Limeaceae Limnanthaceae Limnocharit

aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace

ae Linderniaceae Lindleyaceae Linnaeaceae LippayaceaeLiriaceae Liriodendraceae Li

ssocarpaceae Littorellaceae Loasaceae Lobeliaceaee Loganiaceae LomandraceaeLoni

ceraceae Lopeziaceae Lophiocarpaceae Lophiolaceae Lophiraceae Lophophytaceae L

ophopyxidaceae LoranthaceaeLotaceae Lowiaceae Lupulaceaee Luxemburgiaceae Lu

zuriagaceae Lychnidaceae Lyciaceae LyginiaceaeLygodisodeaceae Lysimachiaceae Ly

thraceae

Macarisiaceae Mackinlayaceae Madiaceae Maesaceae Magnoliaceae Malaceae Male

sherbiaceae MalortieaceaeMalpighiaceae Malvaceae Mangiaceae Manicariaceae Ma

paniaceae Marantaceae Marathraceae MarcgraviaceaeMartyniaceae Mastixiaceae

Matricariaceae Maundiaceae Mayacaceae Medeolaceae Medusagynaceae Peridiscac

eaeMelaleucaceae Melampyraceae Melanophyllaceae Melanthiaceae Melastomatac

eae Meliaceae Melianthaceae MelicaceaeMeliosmaceae Melittidaceae Melochiacea

e Memecylaceae Mendonciaceae Menispermaceae Menthaceae MenyanthaceaeMe

nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

47

Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

Nypaceae Nyssaceae

Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

ceae Papilionaceae Pappophoraceae PapyraceaeParacryphiaceae Parasitaxaceae Par

ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

48

hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

49

cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

50

ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 13: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

13

Heywood et al 2007) may find it of interest to examine a system recognizing only

monophyletic groups it provides a rather different understanding of evolution

Evolutionary classifications in general try and combine phylogeny and

morphological gaps although that is no easy thing to do - it is akin to combining chalk

and cheese (for an attempt to make this impossible task seem to be more objective see

Stuessy amp Koumlnig 2008) However there is a principle from evolutionary classification

that is relevant and I think quite useful here The size of the gap between two groups

tends to be inversely proportional to the sizes of the groups involved (Davis amp Heywood

1963) One can imagine a situation in which a large group is formally divided even

although the distinguishing characters of the two are weak whereas a smaller group

similarly divisable will be left intact

To summarize If hypotheses of phylogeny remain stable we should be able to base

a stable classification on that phylogeny and then get on with our work that is testing

the phylogenies we have elucidating phylogenies in areas where relationships are

unclear studying the evolution of morphology describing species etc In this context

the spread of the Angiosperm Phylogeny Group system (see below) and its widespread

utilisation in technical literature also floras (eg van der Meijden 2005) textbooks (eg

Simpson 2006 Judd et al 2007 [third edition]) dictionaries (Mabberley 2008) more

popular literature (eg Souza amp Lorenzi 2005 Spears 2006) and as an outline for a new

herbarium sequence (Haston et al 2007) is gratifying The posibility that one might be

able to develop a stable phylogeny-based classification of families and in particular

orders represents a dramatic turn-around from the pessimistic attitude about such

higher-level groupings expressed by Davis and Heywood (1963) and Thorne (1976) the

latter even suggesting that we should bury forever the metaphor of the phylogenetic

tree as highly unrealistic (ibid p 56) Returning to Godfray and Knapps (2004) users

of classifications who want a stable informative and accessible classification that

enables easy identification - unfortunately they want cake with everything and cannot

get it - these pages attempt to satisfy as many of their needs as possible but phylogeny

and monophyly are the primary shapers of the APG classification

On this classification in particular

Here I very largely follow the Angiosperm Phylogeny Group classification (APG

2003) Any differences are not to be interpreted as differences in principle simply that

new phylogenies continue to be published and here I attempt to provide an overview

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

14

of current ideas of higher-level relationships of seed plants The Angiosperm Phylogeny

Group classification is based on well-supported relationships evident in the numerous

molecular studies that began to appear in the late 1980s (see APG 1999 for the

principles underlying the classification) but recent phylogenetic work does not

contradict the major outlines of the trees used by APG II (2003) or even those in APG

I (APG 1999) The APG classification has for the most part been conservative and I

am similarly conservative here For the names and authorities I follow APG III (2009)

although the names of the authors may not always be correct these being in something

of a state of flux for the superordinal names used see Chase and Reveal (2009)

although classification at this level here is not exhaustive - only clades with several

orders have superordinal names

Some changes in our ideas of relationships and hence in the clades we talk about

are particularly likely in parts of Lamiales Caryophyllales and Malpighiales for example

although this will not affect the orders themselves Thus some changes to clade

circumscriptions are to be expected although I do not expect them to be substantial

but changes are neither a defect of cladistics nor a necessary consequence of the use of

molecular data Clades are hypotheses of relationships and as hypotheses they may be

overturned However work since APG I has not suggested other than minor changes

in the compositions of the orders even if the odd genus or even family is turning out to

be seriously misplaced - recent examples are Hydatellaceae (from monocots-Poales to

Nymphaeales Saarela et al 2007) Guamatelaceae (from Rosales-Rosaceae to

Crossosomatales Oh amp Potter 2006) and Perrottetia and Bhesa (from Celastrales-

Celastraceae to Huerteales and Malpighiales respectively Zhang amp Simmons 2006) The

main changes have been clarification of the relationhips of individual families or groups

of families that were of uncertain position eg of Chloranthaceae (Moore et al 2007)

Ceratophyllaceae (Jansen et al 2007) and unplaced asterid II families (Winkworth et al

2008a) As mentioned above phylogenies in the area of main interest in these pages are

overwhelmingly tree-like Although there is evidence for acquisition of host plant genes

by parasites (see below) even wider but inexplicable transfer of mitochochondrial

genes in plants like Amborella (Bergthorsson et al 2004 cf Goremykin et al 2009a)

and even transfers of nuclear genes (Vallenback et al 2008) these are the exception

rather than the rule

In cases where the Angiosperm Phylogeny Group II allowed alternatives as to the

limits of families - Papaveraceae in the broad sense or Papaveraceae plus

Pteridophyllaceae plus Fumariaceae Proteaceae in the broad sense or Proteaceae plus

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

15

Platanaceae - the choices made here follow common usage eg as in textbooks like

Judd et al (2007) and Simpson (2006) and particularly in the new edition of

Mabberleys The Plant Book (Mabberley 2008) This latter is itself an attempt to reflect a

consensus the result of taking the opinions of botanists at several meetings A largely

similar consensus is reflected in the APG III classification (APG 2009) where you will

see that alternative classifications have been dispensed with For many people the

existence of such alternative classifications will simply confuse so agreement over

which groupings to use when alternatives are permitted and then dispensing with the

alternatives that are not commonly used seems reasonable I am making the few

changes that are needed to bring the classification here in line with these others but

only gradually since there is not enough time in each day

As already mentioned higher-level relationships in general and the composition of

orders in particular have always presented something of a challenge to systematists

(eg Davis amp Heywood 1963) That being said the composition of some clades like

Apiales Crossosomatales and Pandanales is decidedly unexpected However it is

interesting to see that these higher level clades are generally accepted even in works

with different classificatory philosophies (eg Heywood et al 2007) For clades like

Malpighiales many of the family groupings within Asparagales etc attempts to find

distinctive characters have largely failed (but see Endress amp Matthews 2006a also

the Apomorphies page here) Interestingly as with families some groupings suggested

by molecular studies are supported by morphological andor chemical characters that

have long been known sometimes for over a hundred years the relationship between

Pittosporaceae and ApiaceaeAraliaceae are a case in point (Hegnauer 1969b and

references) As our knowledge of morphology and chemistry improves we can hope for

improvements in the characterisations of clades at all levels

In many cases the new family limits of the Angiosperm Phylogeny Group (see

APG 1999 2003) are not really controversial although changes from the limits

commonly accepted only a decade ago are sometimes quite dramatic (eg Wagenitz

1997) Thus the split of the old Saxifragaceae sl is necessitated by its extreme

polyphyly as also with Icacinaceaesl and Cornaceae sl However the limits and

relationships of such groups had long been considered to be unsatisfactory but until

recently there had been no real reason to prefer one proposed arrangement over

another It is generally accepted that the limits of Lamiaceae and Verbenaceae have to

be redrawn although there is as yet no compelling evidence that the redrawn taxa are

not sister taxa (there is no evidence that they are) But whatever their relationships the

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

16

content of the clades has changed considerably and incidentally they are now easier to

identify than before the decision to recognise two families is not difficult The same is

true for Salicaceae and Achariaceae (Malpighiales) two previously small families that

have received the bulk of the old Flacourtiaceae Clade and hence taxon limits remain

difficult around Theaceae (Ericalessee alsoSladeniaceae Pentaphylacaceae)

and Euphorbiaceae (Malpighiales see

also Phyllanthaceae Picrodendraceae Putranjivaceae etc) Here current groupings

may not represent quite such dramatic changes in our understanding of relationships

Indeed parts of the old Euphorbiaceae that were separated may yet go back together

particularly Phyllanthaceae and Picrodendraceae (see Wurdack et al 2004 Davis et al

2005) even if Theaceae should indeed be dismembered (Geuten et al 2004) However

given that there is no molecular evidence that currently warrants combining all the

segregates of Euphorbiaceae and even if some do come together the clades the

families represent suggest novel groupings not recognised in current classifications

Furthermore Rafflesiaceae appear to be embedded within Euphorbiaceae s str (Davis

et al 2007) so the family is divided Of course maintaining even a moderately narrowly-

circumscribed Euphorbiaceae would entail reducing the iconic Rafflesiaceae to

synonymy Relationships in core Caryophyllales especially

around Phytolaccaceae andNyctaginaceae and also Portulacaceae are incompletely

understood and refashioning of taxon limits will doubtless be needed as cladistic

relationships become apparent (eg see Nyffeler 2007 Ogburn 2007a b) Some

groupings in the old Icacinaceaeand Olacaceae also remain unclear

The discovery of the relationships of parasitic and aquatic groups have presented a

particular challenge to systematists Morphologically some of these plants are so highly

modified that interpretation of the plant body in conventional terms is difficult or even

impossible Thus parasitic groups (broad-leaved angiosperms only) such as Rafflesiaceae

are hard to place since both the vegetative body and the flowers are changed almost

beyond recognition (flowers of rafflesiaceae are a good example of gigantism - Davis et

al 2007) Furthermore plastid gene sequences may be difficult or impossible to obtain

the chloroplast DNA in particular being highly degraded and the rate of molecular

change in general may be high so the problem of long-branch attraction is serious (eg

Duff amp Nickrent 1997 Nickrent et al 1998 Caddick et al 2002a G Petersen et al

2006b) Echlorophyllous sapromycoheterotrophic taxa mostly known from monocots

present similar problems However progress is being made placements for

Apodanthaceae Rafflesiaceae Mitrastemonaceae Cytinaceae and Cynomoriaceae have

recently been suggested (Barkman et al 2004 2007 Davis amp Wurdack 2004 Nickrent et

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

17

al 2004 Davis et al 2007) it seems likely that Burmanniaceae sl are polyphyletic but

both parts are in Dioscoreales (Merckx et al 2006) and relationships within the largely

hemiparasitic Santalales are also gradually being clarified (Maleacutecot 2002) The inclusion

of parasitic taxa in general molecular analyses can cause conniptions (eg Nickrent et al

2004 Davis et al 2004 Chase et al 2006 G Petersen et al 2006b) and there can be

horizontal transmission of genes (eg Davis amp Wurdack 2005 Vitaceae to Rafflesiaceae

Barkman et al 2007 the mitochondrial atp1 gene commonly moves)

Similarly in water plants neither vessels in particular nor much xylem in general is

needed leaves are highly modified and water-mediated pollination if adopted may

well be associated with major changes in floral morphology Here too recent molecular

studies suggest that aquatic groups with hitherto problematic relationships may find

homes Thus Podostemaceae are close to Clusiaceae and Calophyllaceae and are sister

to Hypericaceae (Malpighiales Kita amp Kato 2001) Hydatellaceae which used to be in

Poales are part of Nymphaeales (Saarela et al 2007) and Hydrostachyaceae may be

close to Hydrangeaceae (Cornales Xiang et al 2002) or perhaps should be placed in

Lamiales note that in the first two cases in particular there are morphological and

chemical features that support such a move If Podostemaceae are indeed close to

Clusiaceae I look forward to seeing hypotheses to explain how the dramatic changes in

the vegetative body that have made Podostemaceae so problematic for generations of

systematists took place That conventional wisdom has trouble in understanding or

explaining how the morphologies of groups like Clusiaceae and Podostemaceae can be

related is largely a problem with conventional wisdom Ceratophyllaceae seem to be

finding a position as sister to eudicots (see Moore et al 2007) but their morphology is

so derived that there is no morphological evidence of which I am aware for this

relationship There are also similar although less extreme morphological problems with

many plants pollinated by wind

In such situations especially with some water plants mycoheterotrophs and

parasites a variety of characters may be affected and particularly under such

circumstances groups have tended to be formed using a few or even only a single

character that seems to provide evidence of relationships Add to this the tendency to

weight some characters particularly strongly on a priori grounds and the result was

either the recognition of conglomerate taxa such as Amentiferae which now appear to

be highly polyphyletic or the segregation of families like Plantaginaceae s str (now

much expanded) and Leitneriaceae (now in Simaroubaceae) - all of these are more or

less wind pollinated

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

18

Even although relationships of many parasitic and aquatic groups are being

clarified and at least some of the proposed phylogenetic positions seem to be well

supported it is indeed often still very difficult to understand how such groups have

evolved from their more morphologically conventional relatives However we have had

all sorts of preconceptions as to how characters should evolve Thus we used to

assume that features like highly scalariform vessel perforation plates or the complete

absence of vessels or a flower with an androecium that had many stamens a superior

ovary or separate petals were necessarily plesiomorphic or primitive and conversely

a simple perforation plate and the presence of vessels an androecium with few

stamens inferior ovary or petals that were connnate were almost necessarily

apomorphic or advanced (but cf eg Stebbins 1951) Such assumptions are incorrect

(eg Soltis et al 2005b) Carpels may become secondarily free carpels may fail to close

the seeds then developing outside the confines of the carpel as in some Aspagaraceae-

Nolinoideae Violaceae Berberidaceae Malvaceae-Sterculioideae etc) in Peliosanthes

teta perhaps the only species in Peliosanthes(Asparagaceae-Nolinoideae) the ovary

varies from superior to inferior (Jessop 1976 species limits here need close

investigation see also Kuzoff et al 2001 and Soltis amp Hufford 2002 Saxifragaceae

Apiales etc) many-seeded carpels can evolve from few-seeded carpels

(Razafimandimbison et al 2008) monoecy may be derived from dioecy (Schaefer amp

Renner 2010 and references) Classic studies such as those by Babcock (eg 1947)

on Crepis that assumed that evolution - in this case of the karyotype in particular - was

unidirectional have needed comprehensive re-evaluation (Enke amp Gemeinholzer 2008)

Most if not all characters have reversed andor evolved in parallel as is clear in the

discussion of ovary position here especially in Poales Saxifragales and Asparagales and

also in comments on such morphologically distinctive taxa (when compared with their

immediate relatives) such as Menyanthaceae and especially Pittosporaceae Parallel

evolution may occur even at the level of amino acid substitution as in the independent

acquisition of the phosphoenolpyruvate carboxylase (pepC) gene in C4 photosynthesis in

grasses (Christin et al 2007b see also Blaumlsing et al 2000)

Thinking of how such characters have been used in the past in classifications at the

generic level the results have been similar to those just discussed above for parasitic

plants and others Relying too much on animal pollination syndromes to mark generic

boundaries has all too often led to taxa that are highly unsatisfactory phylogenetically

(see eg Acanthaceae Bignoniaceae Campanulaceae Ericaceae Lamiaceae

Orchidaceae) and over-reliance on characters of fruit and seed (see eg Brassicaceae

and Apiaceae) has also led to unsatisfactory generic limits Again the more general

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

19

problem is the use of one or a very few characters that have been weighted a priori to

structure classifications (see also Garciacutea et al 2009) although it is perhaps not

surprising that such an approach sometimes results in the delineation of taxa that have

indeed turned out to be quite distinct (cf eg Burtt Astragalus versus Oxytropis)

Taxa recognized here are monophyletic yet many are polythetic at the

morphological level that is they lack unique features characterizing (defining) all and

only members of that family they can be recognised phenetically only by the unique

combinations of characters that they posess This is the result of evolution any

synapomorphy characterizing a taxon may be lost or modified beyond easy recognition

in some of its members or the synapomorphy may appear to be identical to a feature

that has evolved in parallel in a quite unrelated plant That plant groups are polythetic is

almost as much a feature of monophyletic taxa as the paraphyletic taxa common in

evolutionary classifications Some families now include substantial variation as

phenetically distinct derived groups are placed in their proper phylogenetic position -

examples are Ericaceae which include the erstwhile Empetraceae a wind-pollinated

group that is florally very different and the overwhelmingly large-flowered and animal-

pollinated Plantaginaceae which now include several derived small-flowered aquatic

and wind-pollinated groups that were previously placed in separate families

Thus some families as delimited here may not be easy to recognize However

remember that detecting relationships - use whatever characters you can even if they

are not obvious - and naming a plant - focus on easy-to-see characters that may not

reflect relationships - are quite different problems Taxa although natural may not be

readily recognizable indeed it was in exactly this context that Lamarck worked out the

basic principles of writing dichotomous keys in 1778 (see Scharf 2007 for keys and the

like) Of course Lamarcks idea of nature was very different from ours - he thought that

there was some kind of continuum of form on which living organisms were to be

situated with no real gaps anywhere - but this meant that his genera (for example)

might well not be sharply distinct from each other even if each were part of the real

continuum that was life Lamarcks separation of the establishment of relationships and

groupings in a system and identification of the plants in that system have been widely

accepted (but cf eg Godfray amp Knapp 2004 many of the contributors to Heywood et

al 2007) Perhaps the best way of identifying plants at the family level is by well-made

multiple access keys as in Watson and Dallwitz (1992a onwards family limits there may

differ substantially from those adopted here) Multiple access keys free users from the

constraints of dichotomous keys in which particular characters are needed at each step

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

20

of the identification process before there can be further progress Instead those

characters that happen to be evident on a specimen can be used in whatever order is

convenient when linked to illustrations glossaries etc their power is enormous (see

Dallwitz et al 2000 [2006] for the principles underlying their construction and use)

Nevertheless dichotomous keys such as those of Hutchinson (1973) and Franz Thonner

(Geesink et al 1981) have their uses Of course taxa coming out adjacent in keys may

well not be at all related

(When identifying large numbers of plants even more efficient than either style of

identification and certainly lots more fun is sight identification Unless you have a

photographic memory you have to build up your knowledge of comparative plant

morphology - on which the ability to make accurate identifications depends - by

repeated observation When faced with an unknown plant I always look for leaf teeth

and stipules distinctive hair types especially stellate and T-shaped hairs glands and

punctations of any sort the presence of latex or other exudate and check leaf insertion

smelling crushed leaves can also be helpful In this context nodal anatomy can usually

be checked using a razor and a hand lens or even simply carefully examining leaf scars

The short paragraphs added after most families may help in confirming familial

identifications)

For the record and for the little that it is worth there are 4 orders and 13 families

of gymnosperms characterised on these pages and together they include some 82

genera and 947 species For angiosperms comparable figures are 56 orders 445

families 13208 genera and 261750 species (of which monocots include 11 orders 89

families 2759 genera and 52760 species) Note however that higher mathematics

was never my strong point and anyway these are pretty meaningless figures even for

species which many (but not all) might concede smacked slightly more of reality than

other taxa estimates range as high as 422000 (Govaerts 2001) Furthermore numbers

of genera and species change daily Nevertheless as emphasized here families are

useful in teaching we as a community can ensure that their limits remain largely stable

and by concentrating on relatively few of them one can gain some familiarity with much

of the worlds flora For summaries of the sizes of orders and families and of the general

arrangement followed in these pages see also the Statistics page itself

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

21

A PHYLOGENETIC TREE OF SEED PLANTS (THE MAIN TREE)

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

22

The Main Tree above is a conservative summary of well supported relationships

between the major clades of extant seed plants Reading - interpreting and

understanding - such a phylogenetic tree is not necessarily easy so reading a tree may

require a bit of practice OHara (eg 1988 1992) provides an accessible introduction to

thinking and talking about history in the context of looking at phylogenetic trees More

recently Krell and Cranston (2004) Crisp and Cook (2005) and others have emphasized

how careful one must be when interpreting and talking about evolution in ladderized

trees in particular and phylogenetic trees in general Gregory (2008) provides a

particularly detailed discussion of this problem In ladderized trees the smaller (in terms

of numbers of terminals) sister taxon is consistently shown on the same side at every

node thus the trees tend to be pectinate like the teeth of a comb however the

horizontal axis that is as a result so evident has no polarity or particular direction The

use of the adjective basal when discussing phylogenetic trees is especially dangerous

(see also Wojciechowski et al 2004 D Soltis et al 2005b) When I use the term and the

context is not otherwise clear I am referring to the pectinations at the base of a

ladderised tree Remember that when talking about sister taxa one can never be basal

to the other or older or younger although members of one can be more derived - in the

sense of having more apomorphies - than the other (but even then one has to be

careful) The words primitive and advanced and lower and higher should

practically never be used especially when talking about taxa Amborellaceae and

Pinaceae are sister to all other flowering plants and Pinales and are basal to the crown

groups of all other flowering plantsPinales respectively but that does not mean they

are thereby primitive Pinaceae in particular have numerous apomorphies The word

plesiomorphic is far less loaded than primitive and can be used to talk about

individual characters

Most trees here have been more or less ladderized as is the one above The

ladderization may be imperfect for example you can see that on the Main Tree

asterids with ten orders follow rosids and relatives which have seventeen although

this is in part because of the currently unresolved nature of relationships between the

main clades of core eudicots (this is likely to change soon) The Main Tree could have

been drawn with Amborella Acorus or a host of other taxa at the far right without

offending any relationships phylogenetic trees are like mobiles the only fixed points

being the nodes Despite the problems just mentioned pectination interpreted

carefully has its value and it is not an arbitrary process As one reads the terminals of a

pectinate tree from left to right adjacent terminals will be separated by apomorphies

that will cumulate in an appropriate fashion When thinking of a book or a herbarium

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

23

sequence (see Haston et al 2007) this is of some value Nymphaeales and

Austrobaileyales are here adjacent on the tree but they could be separated by

hundreds of families in the sequence merely by rotating only the node from which

Nymphaeales and their sister group (all other angiosperms minus about 75 species)

arise If adjacent in a book or herbarium or as in these summary pages then it is

relatively easy to relate their characters both apomorphic and plesiomorphic but if

separated by hundreds of pages or two floors in a large building then it is less easy to

get anything from the sequence (Since all orders in the pages of the main part of this

site are preceded by the apomorphies of all nodes immediately below them in the seed

plant phylogeny and because of the linkages that have been built in to the site this

problem is much less serious there there is no sequence) So the particular sequence of

all seed plants that is based on a phylogenetic classification but is to be used in herbaria

or any other place where taxa have to be arranged linearly as in this part of APweb

will be one that maximizes the number of taxa that are both successive branches of the

tree and placed successively in the sequence Since specimens are generally filed under

families the outline of a new family sequence for arranging herbaria and books can now

be suggested (Haston et al 2007) and such a standardized sequence will further help

teaching and learning about plants

ON THINKING ABOUT APOMORPHIES

Identifying apomorphies is important because understanding the evolution of

morphology in the broad sense - ie including anatomy chemistry etc - in the context

of diversification is one of our major goals For identifying apomorphies several

preconditions must be met One needs to have an accurate well supported phylogeny

one has to have examined the right taxa both from the point of view of morphology and

molecules one has to have coded the characters right (ie delimited states

appropriately) and one has to use an appropriate model of evolution when fitting the

variation to the tree (see eg Omland 1999 Stevens 2006b) If the distributions of

apomorphies below are compared with those in other studies (eg Turgeon et al 2001

Bremer et al 2001 Endress 2001 Albach et al 2001a Judd amp Olmstead 2004 D Soltis

et al 2005b Zhang et al 2006 Judd et al 2007) the reader may find a number of

differences Although I have integrated such studies as far as is possible there are five

reasons that link to the preconditions just mentioned why there may be differences

where features are placed on trees

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

24

1 I may not have found all the information about a particular character there may be

disagreement over its interpretation or I have added information

2 The sampling of nearly all molecular studies is very incomplete (see Salisbury amp Kim

2001 for problems caused by sampling) indeed it is commonplace to decry the

incompleteness of molecular sampling But not only is the sampling in molecular studies

often less than we might wish that of the morphological and chemical characters whose

evolution we are interested in understanding is also often very poor So for many

anatomical chemical and embryological characters that are confidently said to

characterise families and other groups we all too often have no idea if those characters

are applicable to the whole clade or just to a subgroup within that clade However as

our sampling improves we can locate changes on the tree more precisely if in

unexpected positions Thus is is sometimes suggested that Ericaceae have ellagic acid

(Soltis et al 2005b) in fact the little evidence we have indicates that only a small clade

of some 80 species has ellagic acid and the rest of the family - itself alone about one

third of all Ericales - does not Similarly Ericaceae are noted for their

ectendomycorrhizal associations but Enkianthus sister to all other Ericaceae

apparently lacks these (Abe 2005) Thus these mycorrhizae and their associated

distinctive hair roots are unlikely to be apomorphies of Ericaceae - however it appears

that other families near Ericaceae may also have ectendomycorrhizae (see Asai 1934)

3 I am fitting characters to a very conservative tree with many polytomies although the

nodes that are utilised are for the most part strongly supported polytomies make the

optimisation of characters that is the assigment of character state change to a

particular node on the tree notably difficult (eg Madison amp Madison 2002) In nearly

all studies of the evolution of characters distributions of characters are optimised on a

more or less fully resolved tree and the construction of supertrees may yield yet more

detailed hypotheses of relationships (for literature on supertrees see Cotton amp

Wilkinson 2007 2008) Of course some nodes on such fully resolved trees andor

supertrees may have little support and optimisations of characters on such trees may

carry correspondingly little conviction Even parts of some of the trees used here have

poor support eg relationships within aquatic Alismatales etc although I indicate

when this is the case

4 Exactly how one goes about optimising a character on a tree is critically important

Even using simple parsimony optimisations (ACCTRAN or DELTRAN ACCelerated

TRANsitions or DELayed TRANsitions) the position of synapomorphies on trees - and

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

25

hence our ideas of evolution - may differ and this is still more true if one uses maximum

likelihood or Bayesian methods and makes apparently reasonable suggestions about

weighting gains over losses (eg Donoghue amp Ackerley 1996 Cunningham et al 1998

Omland 1997 1999 Ree amp Donoghue 1999 Polly 2001 Webster amp Purvis 2001

Ronquist 2004 Crisp amp Cook 2005) Sannier et al (2007) show how in in Arecaceae that

where on a phylogenetic tree one might peg changes in microsporogenesis will depend

on the methods one uses to do this and Pedersen et al (2007) discuss the sometimes

very substantial effect of node support on the posterior probabilities of ancestral

character states Here I use parsimony optimization not always as explicit as it might be

but I have often indicated where there are particularly important uncertainties as to the

positions of particular character changes on the tree

5 Finally although I have paid quite a lot of attention to the delimitation of the

character states that make up all the characterizations I have not spent enough time on

this critical operation If we are interested in understanding evolution then fitting the

basic variation - not character states - to a tree in principle allows greater flexibility in

understanding morphology in the context of local phylogenies (see also Stevens 2000

Endress 2005c) However many character states used here are delimited globally that

is they are circumscribed in the context of the variation shown by individual characters

across all angiosperms andor in the context of classic ideas of character evolution

Character states often have arbitrary limits and serve best to communicate

information whether they are in fact suitable for either phylogenetic analysis or

understanding evolution are separate issues Studies have rather unsurprisingly

perhaps but importantly shown that dividing the one character into different sets of

states may yield differing ideas of evolution of that character (eg Lamb Frye amp Kron

2003 Hibbett 2004) When looking at trees on which character states are optimised

one should bear in mind the problems surrounding the delimitation of states (eg

Stevens 2000 2006b) and the danger of using pollen or other types - constructs

based on many characters that vary independently but which effectively get lost in

these types thus Blackmore et al (2009) decomposed the pollen types in Asteraceae

into 52 characters

I use Remanes three main criteria of homology or better similarity when

determining the basic similarity of structures on different organisms (see Remane 1952)

These criteria are special properties position and intermediates Special properties

include anatomical or chemical characters gene expression data etc Position refers

to the position of an organ with respect to landmarks on the plant Although plants are

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

26

plants and landmarks are not as fixed as one might like trying to work out the relative

positions of parts is a good way of understanding morphology so long as one does not

try to out-do Procrustes Intermediates include intermediates found on the same

plant or on different plants Intermediates may observed during development or be

adult structures Thus one may find changes during development which suggest what

the nature of a particular structure is and very different stuctures on different plants

may turn out to be similar early in development Or one can simply compare different

taxa and see that structures that appear to be very different when comparing two

different taxa can be linked morphologically by looking at other taxa

Nevertheless the use of these criteria may not yield an unambiguous answer as to

what a structure is even given a solid phylogeny and an improved understanding of

development (see Jaramillo amp Kramer 2007 for a useful discussion) As Endress (2005c)

observed a number of features - position function development shape anatomy

histology gene activity and relationships to other taxa that clearly have petals - can be

used to distinguish a petal (for example) from other floral structures if a petal does not

have one of these features is it thereby not a petal Thus Maturen et al (2005) recently

found that floral organ diversity genes (B and C) were expressed in the large white

inflorescence bracts of Cornus (see also Costa et al 2005) Peney et al (2005) noted that

not all monosulcate pollen grains in monocots have the same developmental pathway

and that as a result such pollen might not have the same ancestral state Reeves and

Olmstead (2003) suggested that the genetic mechanisms causing monosymmetry in

Lamiales and Solanales were different and Serna and Martin (2006) described similar

problems with the development of hairs in Arabidopsis when compared with that of

hairs in Antirrhinum and Solanaceae Indeed as one perhaps might expect delimitation

of states and characters does not necessarily become easier with increasing knowledge

of development etc Thus Buzgo et al (2004) Matthews and Endress (2005) and others

have shown how hard it can be to distinguish between eg prophylls and other floral

structures as their behaviour is studied during the course of the development of the

flower Similarly at what concentration is a particular secondary metabolite deemed to

be present (Waterman (2007)

The validity of the approach used here that of fitting morphological variation to a

largely molecular-based tree may be questioned However I think it rather unlikely that

well-supported molecular branches will be overturned by morphological data Indeed

analyses of morphological data alone do provide support for many of the clades evident

in molecule-only analyses and analyses with morphological and molecular data

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

27

together may lead to increased support for clades (eg Hufford 1992 Nandi et al 1998

[but adding morphological data reduces support for a number of critical clades too]

Doyle amp Endress 2000 however in none of these papers is the use of morphology

without ambiguity) It is unfortunately clear that the use of morphology alone may not

suggest problems in the phylogenetic placement of taxa that later turn out to have been

wrongly included (eg Zhang et al 1992) Thus I have been wary of putting much weight

on clades that have only morphological support but note that I have treated molecule-

based clades with low bootstrap or jacknife support values (esp below 70) or low

posterior probabilies (below 095) likewise Although I may have been mistaken in

placing so much emphasis on molecular data in terms of providing the basic

phylogenetic framework for angiosperms morphological and molecular data are only

very rarely in irreconcilably strong conflict There are indeed a few places where the

conflict seems extreme These include the relative positions of the Monimiaceae and

Hernandiaceae (Laurales) the position of Hanguanaceae (Commelinales [as here] or

Zingiberales) and of Triplostegia (is it in Dipsacaceae or Valerianaceae - see

Dipsacales) Fossils are unlikely to affect the topologies of the trees presented here but

see below for their importance in understanding morphological evolution in general and

the evolution of angiosperms in particular Nevertheless some largely reject the idea

that trees based on molecular data alone can recover phylogenetic relationships

especially when branching points are old and prefer to used trees based on analysis of

morphological data including those taken from fossils (Hilton amp Bateman 2006 Farjon

2007)

All in all however the extent of the congruence between morphological and

molecular data is impressive and heartening and many clades can be characterised

morphologically It seemed in 1998 that there were no unambiguous morphological

synapomorphies for angiosperm orders (K Bremer 2000) and this is still true if by

unambiguous is meant non-homoplasious However many orders and other clades

have synapomorphies even if these may be indistinguishable at least at the current

level of morphological and developmental knowledge from parallel occurences

elsewhere As our morphological knowledge increases so too does the number of

apomorphies

Endress and Matthews (2006a) emphasize the importance of tendencies and

developmental constraints when thinking about characters of clades within the rosids -

many characters or character combinations occur in a rather sporadic fashion within

the clade and apparently notably less frequently outside it There are a number of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

28

examples of tendencies here and in the individual order pages such as the features

enclosed by parentheses in groups above the level of family eg see (cuticular waxes

as aggregated rodlets) for the commelinids and the discussion of the distribution of

polyandry that is flowers with numerous stamens in the asterid I + II groups etc

Tendencies may involve more than single characters When thinking of the

associations of features like integument thickness and vascularization of the integument

(Endress amp Matthews 2006a) one can imagine fairly simple developmental

preconditions being operative As Endress and Matthews (2006a) note it is difficult to

imagine vascular tissue developing in an integument only two cells thick Changes here

would be loosely correlated if morphologically linked However the strongly correlated

changes noted by Givnish et al (2005) are ecologically linked but are presumably

morphologicallydevelopmentally independent When there seem to be characters

evolving more or less together tests can be carried out to see if the changes are

concentrated on certain branches of the tree (eg Maddison 1990 Sanderson 1991

Maddison amp Maddison 2000) Such tests have rarely been carried out at higher levels in

angiosperms

One of the most striking examples of a tendency is the distribution of N-fixation

restricted as it is to a monophyletic group of four clades although it has arisen seven (or

perhaps several more) times independently within the N-fixing clade and several

members of two quite different kinds of bacteria are involved (eg Clawson et al 2004

Elliott et al 2007 Sprent amp James 2007) Other examples of work that bears on the issue

of tendencies include the findings that flowers of polysymmetrical Arabidopsis have

genes like TCP1 that are expressed asymmetrically during early development and TCP1

is a probable orthologue of the well-known CYC gene of Antirrhinum that is involved in

the development of monosymmetric slowers there (Cubas et al 2001 Costa et al 2005

etc) Parallelism might build on this underlying morphologically cryptic monosymmetry

even if details of the genetic mechanisms causing the monosymmetry evident in

particular groups may be different (Reeves amp Olmstead 2003 Cubas 2004 see above)

Similarly there have been several recent suggestions that the capability to synthesise a

particular metabolite may be switched off but not lost and so can sometimes be

reacquired (eg Wink amp Witte 1983 Wink 2003 Liscombe et al 2005 Larsson 2007

Waterman 2007) Hence perhaps the rather spotty distribution of many secondary

metabolites like ellagic acid the indole alkaloid camptothecin iridoids etc when

considered in the context of phylogenies As a non-botanical example - but a rather nice

one - Salwini-Plawen and Mayr (1961) suggested some time ago that there has been

considerable parallelism (40-65 or more independent origins) in the evolution of eyes in

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

29

metazoans However Pax 6 seems to be a master control gene that is involved in eye

formation perhaps throughout all bilateralians (eg Gehring amp Ikeo 1999 Erwin amp

Davidson 2002) which changes the nature of the problem Note however that in

general our invocation of evolutionary tendencies to explain the patterns of

distributions of characters is really an admission that we do not understand those

patterns

As we find out more about variation we will probably find fewer and fewer features

constant throughout a group Most unqualified statements of presence and absence

should properly be qualified as usually present or usually absent if one is thinking of

the characterisations on the order pages as encompassing the total variation within a

clade Thus Pistia alone among monocots as so far known has sieve tube plastids with

starch grains not protein crystals However this does not affect the fact that sieve tube

plastids with cuneate protein crystals and lacking starch grains are an apomorphy for

monocots Furthermore fossils in a number of cases suggest character combinations

unknown in extant taxa as may be seen in the discussions of Fagaceae Platanaceae

Iteaceae Calycanthaceae etc Confusing the issue there may be questions as to where

exactly on the tree a particular fossil is to be placed (eg see Nymphaeaceae

Calycanthaceae Archaefructus etc)

To summarize given our current understandings of both phylogenies and

characters evolution of some characters in which we are interested seems very labile

(see eg D Soltis et al 2005b Endress and Matthews 2006a Stevens 2006b Ekman et

al 2008 for a good recent study) and I have been cautious when talking about character

evolution Much effort must continue to be spent in summarizing characters of well-

established clades at all levels providing features by which they may be recognized and

signaling synapomorphies Remember that (1) the basic morphological anatomical and

chemical knowledge of many critical taxa is woefully incomplete (2) different

assumptions about character evolution may greatly affect the position of

synapomorphies on trees (3) in many cases relationships within and between many

groups are too uncertain at present to worry very much about synapomorphies and

(4) we must be clear about what we do and do not not know As mentioned above I

have indicated in a number of places where there are particularly important

uncertainties as to where characters should be placed on the tree Nevertheless it is a

relatively easy matter to update notes such as these and it can be a simple matter to

incorporate new data on characters that have never before been considered in the

context of a tree Much basic - and unfortunately perhaps unfashionable - work must be

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

30

carried to clarify the distribution of morphological anatomical and chemical characters

for examples of what can to be done see the work of P K Endress and collaborators

(floral morphology and development) and S R Jensen and collaborators (iridoids) But

acquiring information about nodal anatomy cell and tissue distribution nodal anatomy

and the like is not difficult at all and should be generally encouraged

But all this is merely a necesary prelude to the understanding of evolution For this

one needs to know a lot more including dating the phylogeny understanding the

function(s) of characters factoring in relevant aspects of the palaeoenvironment in

which an apomorphy first appeared etc

SUMMARY OF THE SYSTEM

Below is a formalised summary of the relationships within orders of the families of seed

plants There are a few families that are not recognised even as options in APG II and

vice versa as well as a few extra orders The families to be recognised in the most

recent edition of Mabberleys The Plant Book (Mabberley 2008) are also largely

consistent with those below But all differences are trivial and will - I hope - eventually

disappear

Square brackets - [] - enclose clades the plus sign - + - designates sister taxa a

comma - - denotes part of a polytomy and quotation marks - - denotes a

paraphyletic group

SEED PLANTS

GYMNOSPERMS

Cycadales

Cycadaceae + Zamiaceae

Ginkgoales

Ginkgoaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

31

Pinales

Pinaceae [[Araucariaceae [Podocarpaceae + Phyllocladaceae]] [Sciadopityaceae

[Taxaceae + Cupressaceae]]]

Gnetales

Ephedraceae [Gnetaceae + Welwitschiaceae]

ANGIOSPERMSFLOWERING PLANTS

Amborellales

Amborellaceae

Nymphaeales

Hydatellaceae [Cabombaceae + Nymphaeaceae]

Austrobaileyales

Austrobaileyaceae [Schisandraceae + Trimeniaceae]

Chloranthales

Chloranthaceae

MAGNOLIIDS

Magnoliales

Myristicaceae [Magnoliaceae [[Himantandraceae + Degeneriaceae] [Eupomatiaceae +

Annonaceae]]]

Laurales

Calycanthaceae [[Siparunaceae [Gomortegaceae + Atherospermataceae]] [Monimiaceae

[Hernandiaceae + Lauraceae]]]

Canellales

Canellaceae + Winteraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

32

Piperales

[Hydnoraceae + Aristolochiaceae] [Piperaceae + Saururaceae]

MONOCOTS

Acorales

Acoraceae

Alismatales

Araceae [Tofieldiaceae [[Alismataceae [Hydrocharitaceae + Butomaceae]]

[Scheuchzeriaceae [Aponogetonaceae [Juncaginaceae [Maundiaceae [[Posidoniaceae

[Ruppiaceae + Cymodoceaceae]] [Zosteraceae + Potamogetonaceae]]]]]]]

Petrosaviales

Petrosaviaceae

Dioscoreales

Nartheciaceae [[Taccaceae + Thismiaceae] [Burmanniaceae + Dioscoreaceae]]

Pandanales

Velloziaceae Triuridaceae Stemonaceae [Pandanaceae + Cyclanthaceae]

Liliales

Corsiaceae [Campynemataceae [Petermanniaceae [Colchicaceae + Alstroemeriaceae]]

Melanthiaceae [[Philesiaceae + Rhipogonaceae] [Smilacaceae + Liliaceae]]]

Asparagales

Orchidaceae [[Boryaceae [Blandfordiaceae [Lanariaceae [Asteliaceae + Hypoxidaceae]]]]

[[Ixioliriaceae + Tecophilaeaceae] [Doryanthaceae [Iridaceae [Xeronemataceae

[Xanthorrhoeaceae [Alliaceae + Asparagaceae]]]]]]]

COMMELINIDS

Unplaced

Dasypogonaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

33

Arecales

Arecaceae

Poales

[Typhaceae + Bromeliaceae] [Rapateaceae [[Thurniaceae [Juncaceae + Cyperaceae]]

[[Mayacaceae [Eriocaulaceae + Xyridaceae]] [Flagellariaceae [Anarthriaceae

[Centrolepidaceae + Restionaceae]]] [Joinvilleaceae [Ecdeiocoleaceae + Poaceae]]]]]]]

Commelinales

[Commelinaceae + Hanguanaceae] [Philydraceae [Haemodoraceae + Pontederiaceae]]

Zingiberales

Musaceae [Strelitziaceae + Lowiaceae] Heliconiaceae [[Cannaceae + Marantaceae]

[Costaceae + Zingiberaceae]]

Ceratophyllales

Ceratophyllaceae

EUDICOTS

Ranunculales

Eupteleaceae [Papaveraceae [[[Lardizabalaceae + Circaeasteraceae] [Menispermaceae

[Berberidaceae + Ranunculaceae]]]]

Sabiales

Sabiaceae

Proteales

Nelumbonaceae [Platanaceae + Proteaceae]

Trochodendrales

Trochodendraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

34

Buxales

Haptanthaceae [Buxaceae + Didymelaceae]

CORE EUDICOTS

Gunnerales

Gunneraceae + Myrothamnaceae

Dilleniales

Dilleniaceae

Saxifragales

Peridiscaceae [[Paeoniaceae [Altingiaceae [Hamamelidaceae [Cercidiphyllaceae +

Daphniphyllaceae]]]] [[Crassulaceae [Aphanopetalaceae [Tetracarpaeaceae

[Penthoraceae + Haloragaceae]]]] [Iteaceae [Grossulariaceae + Saxifragaceae]]]]

Cynomoriaceae unplaced

Vitales

Vitaceae

ROSIDS

FABIDROSID I

Zygophyllales

Krameriaceae + Zygophyllaceae

Celastrales

Lepidobotryaceae + Celastraceae

Oxalidales

Huaceae [[Connaraceae + Oxalidaceae] [Cunoniaceae [Elaeocarpaceae [Brunelliaceae +

Cephalotaceae]]]]

Malpighiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

35

[Achariaceae [Goupiaceae [Violaceae + Passifloraceae] [Lacistemataceae Salicaceae]]

[Lophopyxidaceae Putranjivaceae] [Ctenolophonaceae [Erythroxylaceae +

Rhizophoraceae]] Linaceae Ixonanthaceae Humiriaceae Irvingiaceae

Centroplacaceae Pandaceae [[Bonnetiaceae + Clusiaceae] [Calophyllaceae

[Hypericaceae + Podostemaceae]]] [Malpighiaceae + Elatinaceae] Ochnaceae

[Peraceae [Rafflesiaceae + Euphorbiaceae]] [Phyllanthaceae + Picrodendraceae]

[Balanopaceae [[Trigoniaceae + Dichapetalaceae] [Chrysobalanaceae +

Euphroniaceae]]] Caryocaraceae

N-FIXING CLADE

Fabales

Quillajaceae [Fabaceae [Polygalaceae + Surianaceae]]

Rosales

Rosaceae [[Barbeyaceae [Elaeagnaceae Dirachmaceae Rhamnaceae]] [Ulmaceae

[Cannabaceae [Moraceae + Urticaceae]]]]

Cucurbitales

Anisophylleaceae [[Corynocarpaceae + Coriariaceae] [Cucurbitaceae [Tetramelaceae

[Datiscaceae + Begoniaceae]]]] Apodanthaceae

Fagales

Nothofagaceae [Fagaceae [[Myricaceae + Juglandaceae] [Casuarinaceae

[Ticodendraceae + Betulaceae]]]]

MALVIDROSID II

pgtGeraniales

Geraniaceae [[Melianthaceae + Francoaceae] [Vivianaceae + Ledocarpaceae]]

Myrtales

Combretaceae [[Onagraceae + Lythraceae] [[Vochysiaceae + Myrtaceae]

[Melastomataceae [Crypteroniaceae [Alzateaceae + Penaeaceae]]]]]

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

36

Crossosomatales

[Staphyleaceae [Guamatelaceae [Crossosomataceae + Stachyuraceae]]] [Aphloiaceae

[Geissolomataceae + Strasburgeriaceae]]

Picramniales

Picramniaceae

Sapindales

Biebersteiniaceae Nitrariaceae [Kirkiaceae [Anacardiaceae + Burseraceae]]

[Sapindaceae [Simaroubaceae Rutaceae Meliaceae]]

Huerteales

Gerradinaceae [Dipentodontaceae + Tapisciaceae]

Malvales

Neuradaceae [Thymelaeaceae [Sphaerosepalaceae Bixaceae [Cistaceae

[Sarcolaenaceae + Dipterocarpaceae]] [Cytinaceae + Muntingiaceae] Malvaceae]]

Brassicales

[Akaniaceae + Tropaeolaceae] [[Moringaceae + Caricaceae] [Setchellanthaceae

[Limnanthaceae [[Koeberliniaceae [Bataceae + Salvadoraceae]] [Emblingiaceae

[Pentadiplandraceae [Gyrostemonaceae Resedaceae Stixis and relatives] Tovariaceae

[Capparaceae [Cleomaceae + Brassicaceae]]]]]]]]

Berberidopsidales

Aextoxicaceae + Berberidopsidaceae

Santalales

Erythropalaceae Olacaceae [[Loranthaceae [Misodendraceae + Schoepfiaceae]]

[Opiliaceae + Santalaceae]] Balanophoraceae unplaced

Caryophyllales

[[Droseraceae [Nepenthaceae [Drosophyllaceae [Ancistrocladaceae +

Dioncophylleaceae]]]]] [[Frankeniaceae + Tamaricaceae] [Polygonaceae +

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

37

Plumbaginaceae]]] [Rhabdodendraceae [Simmondsiaceae [[Asteropeiaceae +

Physenaceae] [[Caryophyllaceae [Achatocarpaceae + Amaranthaceae]]

[Stegnospermataceae [Limeaceae [[Lophocarpaceae [Barbeuiaceae [Aizoaceae

[Sarcobataceae Phytolaccaceae Nyctaginaceae]]]] [Molluginaceae [Halophytaceae

Montiaceae Basellaceae Didiereaceae [Talinaceae [Portulacaceae [Anacampseros etc +

Cactaceae]]]]]]]]]]]]]

ASTERIDS

Cornales

[Cornaceae + Nyssaceae] [Hydrangeaceae + Loasaceae] Hydrostachyaceae [Curtisiaceae

+ Grubbiaceae]

Ericales

[Balsaminaceae [Marcgraviaceae + Tetrameristaceae]] [[Polemoniaceae +

Fouquieraceae] Lecythidaceae [[Sladeniaceae + Pentaphylacaceae] [Sapotaceae

[Ebenaceae [Maesaceae [Theophrastaceae [Myrsinaceae + Primulaceae]]]]]

[Mitrastemonaceae Theaceae [Symplocaceae [Styracaceae + Diapensiaceae]]

[[Sarraceniaceae [Roridulaceae + Actinidiaceae] [Clethraceae [Cyrillaceae +

Ericaceae]]]]]]

LAMIIDASTERID I

Unplaced

Metteniusaceae Oncothecaceae Icacinaceae and some unplaced ex-Icacinaceae

Garryales

Garryaceae + Eucommiaceae

Unplaced

Boraginaceae Vahliaceae

Gentianales

Rubiaceae [Gentianaceae [Loganiaceae [Gelsemiaceae + Apocynaceae]]]

Lamiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

38

Plocospermataceae [Oleaceae [Tetrachondraceae [Calceolariaceae

[Gesneriaceae Peltanthera [Pedaliaceae Carlemanniaceae Martyniaceae Stilbaceae

Plantaginaceae Linderniaceae Bignoniaceae Verbenaceae Lamiaceae Paulowniaceae

Schlegeliaceae Thomandersiaceae Phrymaceae Rehmannia Acanthaceae

Scrophulariaceae Orobanchaceae Byblidaceae Lentibulariaceae]]]]]

Solanales

[Montiniaceae [Sphenocleaceae + Hydroleaceae]] [Convolvulaceae + Solanaceae]

CAMPANULIDASTERID II

Aquifoliales

[[Cardiopteridaceae + Stemonuraceae] [Aquifoliaceae [Helwingiaceae +

Phyllonomaceae]]

Asterales

[Rousseaceae + Campanulaceae] Pentaphragmataceae [[Alseuosmiaceae [Phellinaceae

+ Argophyllaceae]] Stylidiaceae [Menyanthaceae [Goodeniaceae [Calyceraceae +

Asteraceae]]]]

Escalloniales

Escalloniaceae

Bruniales

[Bruniaceae + Columelliaceae]

Apiales

Pennantiaceae [Torricelliaceae [Griseliniaceae [Pittosporaceae [Araliaceae

[Myodocarpaceae + Apiaceae]]]]]

Paracryphiales

Paracryphiaceae

Dipsacales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

39

Adoxaceae [Diervillaceae [Caprifoliaceae [Linnaeaceae [Morinaceae [Dipsacaceae +

Valerianaceae]]]]]

LINKS TO ORDERS AND FAMILIES

[Back to Top]

MAIN GROUPINGS

Asterids commelinids core eudicots asterid 1 asterid

2 eudicots gymnosperms Magnoliophyta monocots N-fixing claderosids seed plants

ALPHABETICAL LISTING OF ALL ORDINAL NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | L | M | N | O | P | Q | R | S | T | U | V | W | X | Z

Abietales Acanthales Acerales Acorales Actinidiales Actinostrobales Adoxales Aescul

ales Agavales Aizoales AkanialesAlismatales Alliales Alseuosmiales Alstroemeriales

Altingiales Amaranthales Amaryllidales Amborellales AmbrosialesAmmiales Amomal

es Ancistrocladales Anisophylleales Annonales Anthobolales Apiales Apocynales Apo

nogetonalesAquifoliales Arales Araliales Aralidiales Araucariales Arecales Aristolochi

ales Athrotaxidales Asarales AsclepiadalesAsparagales Asphodelales Asparagales Ast

eliales Atriplicales Aucubales Austrobaileyales Avenales

Balanitales Balanopales Balanophorales Balsaminales Barbeyales Barclayales Batales

Begoniales BerberidalesBerberidopsidales Betulales Biebersteiniales Bignoniales Bix

ales Boraginales Brassicales Brexiales Bromeliales BrunialesBrunoniales Burmannial

es Burserales Butomales Buxales Byblidales

Cactales Callitrichales Calycanthales Calycerales Campanulales Campynematales Can

ellales Cannales CapparalesCaprifoliales Cardiopteridales Carduales Caricales Carle

manniales Caryophyllales Cassiales Casuarinales CelastralesCentrolepidales Cephalot

ales Cephalotaxales Ceratophyllales Cercidiphyllales Chenopodiales Chironiales Chlor

anthalesChrysobalanales Cinchonales Circaeasterales Cistales Citrales Cocosales Col

chicales Columelliales CombretalesCommelinales Connarales Convolvulales Coriarial

es Cornales Corylales Corynocarpales Crassulales CrossosomatalesCucurbitales Cunn

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

40

inghamiales Cunoniales Cupressales Cycadales Cyclanthales Cymodoceales Cynarales

CynomorialesCyperales Cyrillales Cytinales

Daphnales Daphniphyllales Dasypogonales Datiscales Degeneriales Desfontainiales D

iapensiales Didymelales DillenialesDioales Dioncophyllales Dioscoreales Diospyrales

Dipentodontales Dipsacales Droserales

Ebenales Echiales Elaeagnales Elaeocarpales Elatinales Elodeales Emmotales Empetr

ales Ephedrales Ericales EriocaulalesErythropalaless Escalloniales Eucommiales Euph

orbiales Eupomatiales Eupteleales Euryalaless

Fabales Fagales Falcatifoliales Ficales Flacourtiales Flagellariales Fouquieriales Franc

oales Frangulales

Galiales Garryales Geissolomatales Gentianales Geraniales Gesneriales Ginkgoales G

laucidiales Globulariales GnetalesGoodeniales Greyiales Griseliniales Grossulariales

Grubbiales Gunnerales Gyrocarpales Gyrostemonales

Haemodorales Haloragales Hamamelidales Hanguanales Heisteriales Helleborales He

lwingiales Himantandrales HippuridalesHomaliales Hortensiales Huales Huerteales

Hydatellales Hydnorales Hydrangeales Hydrastidales HydrocharitalesHydropeltidales

Hydrostachyales Hypericales Hypoxidales

Icacinales Illiciales Iridales Irvingales Iteales Ixiales

Jasminales Juglandales Julianiales Juncaginales Juncales

Lacistematales Lactoridales Lamiales Lardizabalales Laurales Lecythidales Ledocarpal

es Leitneriales LentibularialesLigustrales Liliales Limnanthales Linales Liriales Loasal

es Lobeliales Loganiales Lonicerales Loranthales LowialesLythrales

Magnoliales Malpighiales Malvales Marathrales Marcgraviales Mayacales Medusagy

nales Medusandrales MelanthialesMelastomatales Meliales Melianthales Meliosmal

es Menispermales Menyanthales Metteniusales MitrastemonalesMiyoshiales Moni

miales Moringales Musales Myricales Myristicales Myrothamnales Myrsinales Myrta

les

Najadales Nandinales Narcissales Nartheciales Nelumbonales Nepenthales Neuradal

es Nitrariales Nolanales NothofagalesNyctaginales Nymphaeales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

41

Ochnales Oenotherales Olacales Oleales Onagrales Oncothecales Opuntiales Orchid

ales Oxalidales

Saxifragales Pandales Pandanales Papaverales Paracryphiales Parasitaxales Paridales

Parnassiales PassifloralesPenaeales Pennantiales Pentaphragmatales Peridiscales P

etiveriales Petrosaviales Phellinales Philydrales PhyllanthalesPhysenales Phytolaccal

es Picramniales Pinales Pinguiculales Piperales Pittosporales Plantaginales Platanale

sPlumbaginales Poales Podocarpales Podophyllales Podostemales Polemoniales Pol

ygalales Polygonales PontederialesPortulacales Posidoniales Potamogetonales Primu

lales Proteales

Quercales Quillajales Quintiniales

Rafflesiales Ranunculales Rapateales Resedales Restionales Rhabdodendrales Rhamn

ales Rhinanthales RhizophoralesRhodorales Rhoipteleales Roridulales Rosales Rouss

eales Rubiales Ruppiales Rutales

Sabiales Salicales Salvadorales Samolales Samydales Sanguisorbales Santalales Sapin

dales Sapotales SarracenialesSaxegotheales Saxifragales Scheuchzeriales Sciadopitya

les Scleranthales Scrophulariales Scyphostegiales SedalesSilenales Simmondsiales S

milacales Solanales Sphenocleales Sphenostemonales Stangeriales Stellariales Stemo

nalesStilbales Stylidiales Styracales Surianales

Taccales Tamales Tamaricales Taxales Taxodiales Tecophilaeales Terebinthales Tern

stroemiales Theales TheligonalesThymelaeales Tiliales Tofieldiales Torricelliales Tov

ariales Tribelales Trilliales Trimeniales Triuridales TrochodendralesTropaeolales Turn

erales Typhales

Ulmales Urticales

Vacciniales Vahliales Vallisneriales Velloziales Veratrales Verbenales Viburnales Vinc

ales Violales Viscales VitalesVochysiales

Welwitschiales Winterales

Xanthorrhoeales Ximeniales Xyridales

Zamiales Zingiberales Zosterales Zygophyllales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

42

ALPHABETICAL LISTING OF ALL FAMILY NAMES OF SEED PLANTS WITH LINKS

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In some cases clicking on a family synonym will send you directly to the subfamily in

which the genus of the basionym is to be found

Abietaceae Abolbodaceae Abrophyllaceae Acalyphaceae Acanthaceae [Acanthochla

mydaceae] Acarnaceaee AceraceaeAchariaceae Achatocarpaceae Achradaceae Achy

ranthaceae Acmopylaceae Acoraceae Acristaceae ActaeaceaeActinidiaceae Actinota

ceae Amentotaxaceae Adenogrammaceae Adoxaceae Aegialitidaceae Aegicerataceae

Aegilopaceae Aeginetiaceaee Aegiphilaceae Aesculaceae Aextoxicaceae Agapanthac

eae Agavaceae AgdestidaceaeAgialidaceae Agrimoniaceae Agrostidaceae Ailanthace

ae Aitoniaceae Aizoaceae Akaniaceae Alangiaceae AlchemillaceaeAldrovandaceae A

lismataceae Alliaceae Allioniaceae Allophylaceae Aloaceae Alopecuraceae Alpiniacea

e AlseuosmiaceaeAlsinaceae Alsodeiaceae Alstroemeriaceae Altingiaceae Alzateacea

e Amaranthaceae Amaryllidaceae Amborellaceae Ambrosiaceae Amentotaxaceae A

mmanniaceae Ammiaceae Amomaceae Ampelidopsaceae Amygdalaceae Amyridacea

eAnacardiaceae Anagallidaceae Anarthriaceae Anchusaceae Ancistrocladaceae Andr

omedaceae AndropogonaceaeAndrostachyaceae Androsynaceae Anemarrhenaceae

Anemonaceae Angelicaceae Anisophylleaceae AnnonaceaeAnomochloaceae Anopter

aceae Anrederaceae Anthemidaceae Anthericaceae Anthobolaceae Antidesmataceae

Antirrhinaceae Antoniaceae Aparinaceae Aphanopetalaceae Aphloiaceae Aphyllanth

aceae Apiaceae ApocynaceaeApodanthaceae Aponogetonaceae Aporusaceae Apose

ridaceae Apostasiaceae Aptandraceae AquifoliaceaeAquilariaceae Aquilegiaceae A

raceae Aragoaceae Araliaceae Aralidiaceaee Araucariaceae Arbutaceae Arceuthidace

aeArceuthobiaceae Arctostaphylaceae Arctotidaceae Ardisiaceae Arecaceae Argoph

yllaceae Arisaraceae AristolochiaceaeAristoteliaceae Arjonaceae Armeriaceae Artem

isiaceae Arthrotaxidaceae Artocarpaceae Arundinaceae ArundinellaceaeAsaraceae A

sclepiadaceae Ascyraceaee Asiraceae Asparagaceae Asperulaceae Asphodelaceae As

pidistraceaeeAsteliaceae Asteraceae Asteranthaceae Asteropeiaceae Astragalaceae

Astrocarpaceae AthanasiaceaeAtherospermataceae Atriplicaceae Atropaceae Aucuba

ceae Aurantiaceae Austrobaileyaceae Austrotaxaceae AvenaceaeAverrhoaceae Avet

raceae Avicenniaceae Azaleaceae Azimaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Balanitaceae Balanopaceae Balanophoraceae Balsameaceae Balsaminaceae Bambusa

ceae Barbaceniaceae BarbeuiaceaeBarbeyaceae Barclayaceae Barringtoniaceae Base

llaceae Bataceae Baueraceae Bauhiniaceae Baxteriaceae BegoniaceaeBehniaceae B

elangeraceae Belloniaceae Bembiciaceae Berberidaceae Berberidopsidaceae Berryac

eaee BersamaceaeBertyaceae Berzeliaceae Besleriaceae Betaceae Betulaceae Biebe

rsteiniaceae Bifariaceae Bignoniaceae BischofiaceaeBixaceae Blakeaceae Blandfordia

ceae Blattiaceae Blepharocaryaceae Blitaceae Blyxaceae Boerlagellaceae Bolivaracea

eBombacaceae Bonnetiaceae Bontiaceae Boopidaceae Boraginaceae Borassaceae B

oroniaceae BoryaceaeBotryodendraceae Bougainvilleaceae Boweniaceae Brassicacea

e Brachycaulaceae Bretschneideraceae BrexiaceaeBromeliaceae Brownlowiaceae Br

unelliaceae Bruniaceae Brunoniaceae Brunsvigiaceae Bryoniaceae BuchneraceaeBuci

daceae Buddlejaceae Buglossaceae Bulbocodiaceae Bumeliaceae Bupleuraceae Burc

hardiaceae BurmanniaceaeBurseraceae Butneriaceae Butomaceae Buxaceae Byblida

ceae Byttneriaceae

Cabombaceae Cacaoaceae Cactaceae Caesalpiniaceae Caladiaceae Calamaceae Calce

olariaceae CalectasiaceaeCalendulaceae Callaceae Callicomaceae Calligonaceae Calli

traceae Callitrichaceae Calochortaceae CalophyllaceaeCalthaceae Calycanthaceae Ca

lyceraceae Cambogiaceae Camelliaceae Campanulaceae CampynemataceaeCanacom

yricaceae Canellaceae Cannabaceae Cannaceae Canopodaceae Canotiaceae Cansjera

ceae CapparaceaeCaprariaceae Caprifoliaceae Cardiopteridaceae Carduaceae Carica

ceae Carissaceae Carlemanniaceae CarpinaceaeCarpodetaceae Cartonemataceae Ca

ryocaraceae Caryophyllaceae Casearieaceae Cassiaceae CassipoureaceaeCassythacea

e Castaneaceae Castelaceae Casuarinaceae Catesbaeaceae Cathedraceae Cecropiace

ae CedraceaeCedrelaceae Celastraceae Celosiaceae Celtidaceae Centaureaceae Cen

trolepidaceae Centroplacaceae CepaceaeCephalanthaceae Cephalotaceae Cephalota

xaceae Cerastiaceae Ceratoniaceae Ceratophyllaceae CerberaceaeCercidiphyllaceae

Cercocarpaceae Cercodiaceae Cereaceae Cerinthaceae Ceroxylaceae Cestraceae Cev

alliaceaeChailletiaceae Chamaedoraceae Chamaemoraceae Chamelauciaceae Chaun

ochitonaceae Chelidoniaceae ChelonaceaeChenopodiaceae Chimonanthaceae Chingi

thamnaceae Chionographidaceae Chiranthodendraceae ChironiaceaeChlaenaceae Ch

loanthaceae Chloranthaceae Chloridaceae Chlorogalaceae Chrysobalanaceae Cicerac

eae CichoriaceaeCimicifugaceae Cinchoniaceae Circaeaceae Circaeasteraceae Cissac

eae Cistaceae Citraceae Clematidaceae CleomaceaeClethraceae Clusiaceae Cneorac

eae Cnestidaceae Cnicaceae Cobaeaceae Cochlospermaceae Cocosaceae Coffeaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Colchicaceae Coleogynaceae Columelliaceae Combretaceae Commelinaceae Comocl

adiaceae Compositae CompsoaceaeConantheraceae Connaraceae Conostylidaceae C

onvallariaceae Convolvulaceae Cordiaceae CoreopsidaceaeCoriandraceae Coriariacea

e Coridaceae Corispermaceae Cornaceae Corokiaceae Coronillaceae Correaceae Cor

rigiolaceaeCorsiaceae Corydalaceae Corylaceae Corynocarpaceae Coryphaceae Cost

aceae Cotyledonaceae CoulaceaeCoutariaceae Coutoubiaceae Crassulaceae Crescen

tiaceae Cressaceae Crinaceae Crocaceae CroomiaceaeCrossosomataceae Crotonacea

e Cruciferae Cryptaceae Crypteroniaceae Cryptocorynaceae CryptomeriaceaeCtenol

ophonaceae Cucurbitaceae Cunninghamiaceae Cunoniaceae Cupressaceae Curcumac

eae Curtisiaceae CuscutaceaeCyananthaceae Cyanastraceae Cynanchaceae Cyanella

ceae Cycadaceae Cyclanthaceae CyclantheraceaeCyclocheilaceae Cydoniaceae Cymo

doceaceae Cynaraceae Cynocrambaceae Cynomoriaceae Cyperaceae CyphiaceaeCyp

hocarpaceae Cypripediaceae Cyrillaceae Cyrtandraceae Cyrtanthaceae Cytinaceae

Dacrycarpaceae Dactylanthaceae Damasoniaceae Daphnaceae Daphniphyllaceae Das

ypogonaceae DatiscaceaeDaturaceae Daucaceae Davidiaceae Davidsoniaceae Decais

neaceae Deeringiaceae Degeneriaceae DelphiniaceaeDendrophthoaceae Columelliac

eae Detariaceae Dialypetalanthaceae Dianellaceae Dianthaceae DiapensiaceaeDicha

petalaceae Dichondraceae Diclidantheraceae Dicrastylidiaceae Dictamnaceae Didiere

aceae DidymelaceaeDidymocarpaceae Diegodendraceae Diervillaceae Digitalidaceae

e Dilatridaceae Dilleniaceae Dioaceae DionaeaceaeDioncophyllaceae Dioscoreaceae

Diosmaceae Diospyraceae Dipentodontaceae Diphylleiaceae Diplolaenaceae Dipsaca

ceaeDipterocarpaceae Dirachmaceae Disanthaceae Diselmaceae Dodonaeaceae Do

mbeyaceae Donatiaceae DorsteniaceaeDortmannaceae Doryanthaceae Drabaceae D

racaenaceae Dracontiaceae Drimyidaceae Droseraceae DrosophyllaceaeDryadaceae

Duabangaceae Duckeodendraceae Dulongiaceae Durantaceae Durionaceae Dysphani

aceae

Ebenaceae Eccremidaceae Ecdeiocoleaceae Echiaceae Echinopaceae Ehretiaceae Ela

eagnaceae ElaeocarpaceaeElegiaceae Elatinaceae Ellisiaceae Ellisiophyllaceae Elodea

ceae Elytranthaceae Embeliaceae Emblingiaceae EmmotaceaeEmpetraceae Encepha

lartaceae Engelhardtiaceae Enhalaceae Epacridaceae Ephedraceae Ephemeranthacea

eEpilobiaceae Epimediaceae Eragrostidaceae Eremolepidaceae Eremosynaceae Erica

ceae Erinaceae EriocaulaceaeEriogonaceae Eriospermaceae Erodiaceae Erycibaceae

Eryngiaceae Erysimaceae Erythroniaceae ErythropalaceaeErythrospermaceae Erythro

xylaceae Escalloniaceae Eschscholziaceae Eucomidaceae Eucommiaceae Eucryphiace

aeEuonymaceae Eupatoriaceae Euphorbiaceae Euphrasiaceae Euphroniaceae Eupo

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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matiaceae Eupteleaceae EuryalaceaeEustrephaceae Euthemidaceae Eutocaceae Exb

ucklandiaceae Exocarpaceae

Fabaceae Fagaceae Falcatifoliaceae Ferulaceae Festucaceae Ficaceae Ficoidaceae Fit

zroyaceae FlacourtiaceaeFlagellariaceae Flindersiaceae Foetidiaceae Forestieraceae

Fothergillaceae Fouquieriaceae Fragariaceae FrancoaceaeFrangulaceae Frankeniacea

e Fraxinaceae Fraxinellaceae Freycinetiaceae Fritillariaceae Fuchsiaceae Fumariacea

eFunkiaceae

Gaiadendraceae Galacaceae Galanthaceae Galaxiaceae Galedupaceae Galeniaceae G

aliaceae Garciniaceae GardeniaceaeGardneriaceae Garryaceae Geissolomataceae Ge

itonoplesiaceae Gelsemiaceae Geniostomaceae GentianaceaeGeonomataceae Geosir

idaceae Geraniaceae Gerrardinaceae Gesneriaceae Gethyllidaceae Gilliesiaceae Gina

lloaceaeGinkgoaceae Gisekiaceae Gladiolaceae Glaucidiaceae Glechomaceae Glinace

ae Globulariaceae Gnaphaliaceae GnetaceaeGoetzeaceae Gomortegaceae Gomphiac

eae Gomphrenaceae Gonystylaceae Goodeniaceae Gordoniaceae GouaniaceaeGoup

iaceae Gramineae Gratiolaceae Grewiaceae Greyiaceae Grielaceae Griseliniaceae Gr

onoviaceae GrossulariaceaeGrubbiaceae Guaiacanaceae Guamatelaceae Guettardac

eae Gunneraceae Gustaviaceae Guttiferae GyrocarpaceaeGyrostemonaceae

Hachetteaceae Haemanthaceae Haemodoraceae Halesiaceae Hallieraceae Halocarpa

ceae Halophilaceae HalophytaceaeHaloragaceae Hamamelidaceae Hanguanaceae H

aptanthaceae Harmandaceae Hebenstretiaceae HectorellaceaeHederaceae Hedyosm

aceae Hedyotidaceae Hedysaraceae Heisteriaceae Heleniaceae Heliamphoraceae H

elianthaceaeHelianthemaceae Helichrysaceae Heliconiaceae Helicteraceae Heliotropi

aceae Helleboraceae Heloniadaceae HelosidaceaeHelwingiaceae Hemerocallidaceae

Hemimeridaceae Hemiphylacaceae Henriqueziaceae Henslowiaceae Hermanniaceae

Hernandiaceae Herniariaceae Herreriaceae Hesperocallidaceae Heterantheraceae He

teropyxidaceae HeterostylaceaeHibbertiaceae Hibiscaceae Hilleriaceae Himantandra

ceae Hippocastanaceae Hippocrateaceae HippomanaceaeHippophaeaceae Hippurida

ceae Hirtellaceae Holacanthaceae Homaliaceae Hopkinsiaceae Hoplestigmataceae H

ordeaceaeHornschuchiaceae Hortensiaceae Hortoniaceae Hostaceae Houstoniaceae

Huaceae Huerteaceae HugoniaceaeHumbertiaceae Humiriaceae Hyacinthaceae Hyd

atellaceae Hydnoraceae Hydrangeaceae Hydrastidaceae HydrillaceaeHydrocharitacea

e Hydrocotylaceae Hydrogetonaceae Hydroleaceae Hydropeltidaceae Hydrophylacac

eae HydrophyllaceaeHydrostachyaceae Hymenocardiaceae Hypecoaceae Hyperanth

eraceae Hypericaceae Hypopityaceae HypoxidaceaeHypseocharitaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta

ceae ImperatoriaceaeInocarpaceae Inulaceae Ionidiaceae Iriarteaceae Iridaceae Irvi

ngiaceae Isnardiaceae Isophysidaceae IteaceaeStrasburgeriaceae Ixiaceae Ixioliriace

ae Ixonanthaceae

Jalapaceae Jamboliferaceae Japonoliriaceae Jasionaceae Jasminiaceae Johnsoniaceae

Joinvilleaceae JuglandaceaeJulianiaceae Juncaceae Juncaginaceae Juniperaceae Jus

siaeaceae Justiciaceae

Kadsuraceae Kaliphoraceae Kalmiaceae Kaniaceae Kiggelariaceae Kingdoniaceae Kin

giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae

Krameriaceae

Labiatae Lacandoniaceae Lachenaliaceae Lacistemataceae Lactoridaceae Lactucaceae

Lagerstroemiaceaee LamiaceaeLampsanaceae Lanariaceae Langsdorffiaceae Lantan

aceae Lapageriaceae Lardizabalaceae Lasiopetalaceae LauraceaeLawsoniaceae Laxm

anniaceae Lecythidaceae Ledaceae Ledocarpaceae Leeaceae Leguminosae Leitneriac

eae LemnaceaeLennoaceae Lentibulariaceae Lentiscaceae Leoniaceae Leonticaceae

Lepidobotryaceae LepidocarpaceaeLepidocaryaceae Lepidocerataceae Leptaulaceae

Leptospermaceae Lepturaceae Lepuropetalaceae LeuctenbergiaceaeLibocedraceae

Licaniaceae Ligustraceae Lilaeaceae Liliaceae Limeaceae Limnanthaceae Limnocharit

aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace

ae Linderniaceae Lindleyaceae Linnaeaceae LippayaceaeLiriaceae Liriodendraceae Li

ssocarpaceae Littorellaceae Loasaceae Lobeliaceaee Loganiaceae LomandraceaeLoni

ceraceae Lopeziaceae Lophiocarpaceae Lophiolaceae Lophiraceae Lophophytaceae L

ophopyxidaceae LoranthaceaeLotaceae Lowiaceae Lupulaceaee Luxemburgiaceae Lu

zuriagaceae Lychnidaceae Lyciaceae LyginiaceaeLygodisodeaceae Lysimachiaceae Ly

thraceae

Macarisiaceae Mackinlayaceae Madiaceae Maesaceae Magnoliaceae Malaceae Male

sherbiaceae MalortieaceaeMalpighiaceae Malvaceae Mangiaceae Manicariaceae Ma

paniaceae Marantaceae Marathraceae MarcgraviaceaeMartyniaceae Mastixiaceae

Matricariaceae Maundiaceae Mayacaceae Medeolaceae Medusagynaceae Peridiscac

eaeMelaleucaceae Melampyraceae Melanophyllaceae Melanthiaceae Melastomatac

eae Meliaceae Melianthaceae MelicaceaeMeliosmaceae Melittidaceae Melochiacea

e Memecylaceae Mendonciaceae Menispermaceae Menthaceae MenyanthaceaeMe

nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

Nypaceae Nyssaceae

Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

ceae Papilionaceae Pappophoraceae PapyraceaeParacryphiaceae Parasitaxaceae Par

ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

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hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

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cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

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ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 14: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

14

of current ideas of higher-level relationships of seed plants The Angiosperm Phylogeny

Group classification is based on well-supported relationships evident in the numerous

molecular studies that began to appear in the late 1980s (see APG 1999 for the

principles underlying the classification) but recent phylogenetic work does not

contradict the major outlines of the trees used by APG II (2003) or even those in APG

I (APG 1999) The APG classification has for the most part been conservative and I

am similarly conservative here For the names and authorities I follow APG III (2009)

although the names of the authors may not always be correct these being in something

of a state of flux for the superordinal names used see Chase and Reveal (2009)

although classification at this level here is not exhaustive - only clades with several

orders have superordinal names

Some changes in our ideas of relationships and hence in the clades we talk about

are particularly likely in parts of Lamiales Caryophyllales and Malpighiales for example

although this will not affect the orders themselves Thus some changes to clade

circumscriptions are to be expected although I do not expect them to be substantial

but changes are neither a defect of cladistics nor a necessary consequence of the use of

molecular data Clades are hypotheses of relationships and as hypotheses they may be

overturned However work since APG I has not suggested other than minor changes

in the compositions of the orders even if the odd genus or even family is turning out to

be seriously misplaced - recent examples are Hydatellaceae (from monocots-Poales to

Nymphaeales Saarela et al 2007) Guamatelaceae (from Rosales-Rosaceae to

Crossosomatales Oh amp Potter 2006) and Perrottetia and Bhesa (from Celastrales-

Celastraceae to Huerteales and Malpighiales respectively Zhang amp Simmons 2006) The

main changes have been clarification of the relationhips of individual families or groups

of families that were of uncertain position eg of Chloranthaceae (Moore et al 2007)

Ceratophyllaceae (Jansen et al 2007) and unplaced asterid II families (Winkworth et al

2008a) As mentioned above phylogenies in the area of main interest in these pages are

overwhelmingly tree-like Although there is evidence for acquisition of host plant genes

by parasites (see below) even wider but inexplicable transfer of mitochochondrial

genes in plants like Amborella (Bergthorsson et al 2004 cf Goremykin et al 2009a)

and even transfers of nuclear genes (Vallenback et al 2008) these are the exception

rather than the rule

In cases where the Angiosperm Phylogeny Group II allowed alternatives as to the

limits of families - Papaveraceae in the broad sense or Papaveraceae plus

Pteridophyllaceae plus Fumariaceae Proteaceae in the broad sense or Proteaceae plus

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

15

Platanaceae - the choices made here follow common usage eg as in textbooks like

Judd et al (2007) and Simpson (2006) and particularly in the new edition of

Mabberleys The Plant Book (Mabberley 2008) This latter is itself an attempt to reflect a

consensus the result of taking the opinions of botanists at several meetings A largely

similar consensus is reflected in the APG III classification (APG 2009) where you will

see that alternative classifications have been dispensed with For many people the

existence of such alternative classifications will simply confuse so agreement over

which groupings to use when alternatives are permitted and then dispensing with the

alternatives that are not commonly used seems reasonable I am making the few

changes that are needed to bring the classification here in line with these others but

only gradually since there is not enough time in each day

As already mentioned higher-level relationships in general and the composition of

orders in particular have always presented something of a challenge to systematists

(eg Davis amp Heywood 1963) That being said the composition of some clades like

Apiales Crossosomatales and Pandanales is decidedly unexpected However it is

interesting to see that these higher level clades are generally accepted even in works

with different classificatory philosophies (eg Heywood et al 2007) For clades like

Malpighiales many of the family groupings within Asparagales etc attempts to find

distinctive characters have largely failed (but see Endress amp Matthews 2006a also

the Apomorphies page here) Interestingly as with families some groupings suggested

by molecular studies are supported by morphological andor chemical characters that

have long been known sometimes for over a hundred years the relationship between

Pittosporaceae and ApiaceaeAraliaceae are a case in point (Hegnauer 1969b and

references) As our knowledge of morphology and chemistry improves we can hope for

improvements in the characterisations of clades at all levels

In many cases the new family limits of the Angiosperm Phylogeny Group (see

APG 1999 2003) are not really controversial although changes from the limits

commonly accepted only a decade ago are sometimes quite dramatic (eg Wagenitz

1997) Thus the split of the old Saxifragaceae sl is necessitated by its extreme

polyphyly as also with Icacinaceaesl and Cornaceae sl However the limits and

relationships of such groups had long been considered to be unsatisfactory but until

recently there had been no real reason to prefer one proposed arrangement over

another It is generally accepted that the limits of Lamiaceae and Verbenaceae have to

be redrawn although there is as yet no compelling evidence that the redrawn taxa are

not sister taxa (there is no evidence that they are) But whatever their relationships the

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

16

content of the clades has changed considerably and incidentally they are now easier to

identify than before the decision to recognise two families is not difficult The same is

true for Salicaceae and Achariaceae (Malpighiales) two previously small families that

have received the bulk of the old Flacourtiaceae Clade and hence taxon limits remain

difficult around Theaceae (Ericalessee alsoSladeniaceae Pentaphylacaceae)

and Euphorbiaceae (Malpighiales see

also Phyllanthaceae Picrodendraceae Putranjivaceae etc) Here current groupings

may not represent quite such dramatic changes in our understanding of relationships

Indeed parts of the old Euphorbiaceae that were separated may yet go back together

particularly Phyllanthaceae and Picrodendraceae (see Wurdack et al 2004 Davis et al

2005) even if Theaceae should indeed be dismembered (Geuten et al 2004) However

given that there is no molecular evidence that currently warrants combining all the

segregates of Euphorbiaceae and even if some do come together the clades the

families represent suggest novel groupings not recognised in current classifications

Furthermore Rafflesiaceae appear to be embedded within Euphorbiaceae s str (Davis

et al 2007) so the family is divided Of course maintaining even a moderately narrowly-

circumscribed Euphorbiaceae would entail reducing the iconic Rafflesiaceae to

synonymy Relationships in core Caryophyllales especially

around Phytolaccaceae andNyctaginaceae and also Portulacaceae are incompletely

understood and refashioning of taxon limits will doubtless be needed as cladistic

relationships become apparent (eg see Nyffeler 2007 Ogburn 2007a b) Some

groupings in the old Icacinaceaeand Olacaceae also remain unclear

The discovery of the relationships of parasitic and aquatic groups have presented a

particular challenge to systematists Morphologically some of these plants are so highly

modified that interpretation of the plant body in conventional terms is difficult or even

impossible Thus parasitic groups (broad-leaved angiosperms only) such as Rafflesiaceae

are hard to place since both the vegetative body and the flowers are changed almost

beyond recognition (flowers of rafflesiaceae are a good example of gigantism - Davis et

al 2007) Furthermore plastid gene sequences may be difficult or impossible to obtain

the chloroplast DNA in particular being highly degraded and the rate of molecular

change in general may be high so the problem of long-branch attraction is serious (eg

Duff amp Nickrent 1997 Nickrent et al 1998 Caddick et al 2002a G Petersen et al

2006b) Echlorophyllous sapromycoheterotrophic taxa mostly known from monocots

present similar problems However progress is being made placements for

Apodanthaceae Rafflesiaceae Mitrastemonaceae Cytinaceae and Cynomoriaceae have

recently been suggested (Barkman et al 2004 2007 Davis amp Wurdack 2004 Nickrent et

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

17

al 2004 Davis et al 2007) it seems likely that Burmanniaceae sl are polyphyletic but

both parts are in Dioscoreales (Merckx et al 2006) and relationships within the largely

hemiparasitic Santalales are also gradually being clarified (Maleacutecot 2002) The inclusion

of parasitic taxa in general molecular analyses can cause conniptions (eg Nickrent et al

2004 Davis et al 2004 Chase et al 2006 G Petersen et al 2006b) and there can be

horizontal transmission of genes (eg Davis amp Wurdack 2005 Vitaceae to Rafflesiaceae

Barkman et al 2007 the mitochondrial atp1 gene commonly moves)

Similarly in water plants neither vessels in particular nor much xylem in general is

needed leaves are highly modified and water-mediated pollination if adopted may

well be associated with major changes in floral morphology Here too recent molecular

studies suggest that aquatic groups with hitherto problematic relationships may find

homes Thus Podostemaceae are close to Clusiaceae and Calophyllaceae and are sister

to Hypericaceae (Malpighiales Kita amp Kato 2001) Hydatellaceae which used to be in

Poales are part of Nymphaeales (Saarela et al 2007) and Hydrostachyaceae may be

close to Hydrangeaceae (Cornales Xiang et al 2002) or perhaps should be placed in

Lamiales note that in the first two cases in particular there are morphological and

chemical features that support such a move If Podostemaceae are indeed close to

Clusiaceae I look forward to seeing hypotheses to explain how the dramatic changes in

the vegetative body that have made Podostemaceae so problematic for generations of

systematists took place That conventional wisdom has trouble in understanding or

explaining how the morphologies of groups like Clusiaceae and Podostemaceae can be

related is largely a problem with conventional wisdom Ceratophyllaceae seem to be

finding a position as sister to eudicots (see Moore et al 2007) but their morphology is

so derived that there is no morphological evidence of which I am aware for this

relationship There are also similar although less extreme morphological problems with

many plants pollinated by wind

In such situations especially with some water plants mycoheterotrophs and

parasites a variety of characters may be affected and particularly under such

circumstances groups have tended to be formed using a few or even only a single

character that seems to provide evidence of relationships Add to this the tendency to

weight some characters particularly strongly on a priori grounds and the result was

either the recognition of conglomerate taxa such as Amentiferae which now appear to

be highly polyphyletic or the segregation of families like Plantaginaceae s str (now

much expanded) and Leitneriaceae (now in Simaroubaceae) - all of these are more or

less wind pollinated

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

18

Even although relationships of many parasitic and aquatic groups are being

clarified and at least some of the proposed phylogenetic positions seem to be well

supported it is indeed often still very difficult to understand how such groups have

evolved from their more morphologically conventional relatives However we have had

all sorts of preconceptions as to how characters should evolve Thus we used to

assume that features like highly scalariform vessel perforation plates or the complete

absence of vessels or a flower with an androecium that had many stamens a superior

ovary or separate petals were necessarily plesiomorphic or primitive and conversely

a simple perforation plate and the presence of vessels an androecium with few

stamens inferior ovary or petals that were connnate were almost necessarily

apomorphic or advanced (but cf eg Stebbins 1951) Such assumptions are incorrect

(eg Soltis et al 2005b) Carpels may become secondarily free carpels may fail to close

the seeds then developing outside the confines of the carpel as in some Aspagaraceae-

Nolinoideae Violaceae Berberidaceae Malvaceae-Sterculioideae etc) in Peliosanthes

teta perhaps the only species in Peliosanthes(Asparagaceae-Nolinoideae) the ovary

varies from superior to inferior (Jessop 1976 species limits here need close

investigation see also Kuzoff et al 2001 and Soltis amp Hufford 2002 Saxifragaceae

Apiales etc) many-seeded carpels can evolve from few-seeded carpels

(Razafimandimbison et al 2008) monoecy may be derived from dioecy (Schaefer amp

Renner 2010 and references) Classic studies such as those by Babcock (eg 1947)

on Crepis that assumed that evolution - in this case of the karyotype in particular - was

unidirectional have needed comprehensive re-evaluation (Enke amp Gemeinholzer 2008)

Most if not all characters have reversed andor evolved in parallel as is clear in the

discussion of ovary position here especially in Poales Saxifragales and Asparagales and

also in comments on such morphologically distinctive taxa (when compared with their

immediate relatives) such as Menyanthaceae and especially Pittosporaceae Parallel

evolution may occur even at the level of amino acid substitution as in the independent

acquisition of the phosphoenolpyruvate carboxylase (pepC) gene in C4 photosynthesis in

grasses (Christin et al 2007b see also Blaumlsing et al 2000)

Thinking of how such characters have been used in the past in classifications at the

generic level the results have been similar to those just discussed above for parasitic

plants and others Relying too much on animal pollination syndromes to mark generic

boundaries has all too often led to taxa that are highly unsatisfactory phylogenetically

(see eg Acanthaceae Bignoniaceae Campanulaceae Ericaceae Lamiaceae

Orchidaceae) and over-reliance on characters of fruit and seed (see eg Brassicaceae

and Apiaceae) has also led to unsatisfactory generic limits Again the more general

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

19

problem is the use of one or a very few characters that have been weighted a priori to

structure classifications (see also Garciacutea et al 2009) although it is perhaps not

surprising that such an approach sometimes results in the delineation of taxa that have

indeed turned out to be quite distinct (cf eg Burtt Astragalus versus Oxytropis)

Taxa recognized here are monophyletic yet many are polythetic at the

morphological level that is they lack unique features characterizing (defining) all and

only members of that family they can be recognised phenetically only by the unique

combinations of characters that they posess This is the result of evolution any

synapomorphy characterizing a taxon may be lost or modified beyond easy recognition

in some of its members or the synapomorphy may appear to be identical to a feature

that has evolved in parallel in a quite unrelated plant That plant groups are polythetic is

almost as much a feature of monophyletic taxa as the paraphyletic taxa common in

evolutionary classifications Some families now include substantial variation as

phenetically distinct derived groups are placed in their proper phylogenetic position -

examples are Ericaceae which include the erstwhile Empetraceae a wind-pollinated

group that is florally very different and the overwhelmingly large-flowered and animal-

pollinated Plantaginaceae which now include several derived small-flowered aquatic

and wind-pollinated groups that were previously placed in separate families

Thus some families as delimited here may not be easy to recognize However

remember that detecting relationships - use whatever characters you can even if they

are not obvious - and naming a plant - focus on easy-to-see characters that may not

reflect relationships - are quite different problems Taxa although natural may not be

readily recognizable indeed it was in exactly this context that Lamarck worked out the

basic principles of writing dichotomous keys in 1778 (see Scharf 2007 for keys and the

like) Of course Lamarcks idea of nature was very different from ours - he thought that

there was some kind of continuum of form on which living organisms were to be

situated with no real gaps anywhere - but this meant that his genera (for example)

might well not be sharply distinct from each other even if each were part of the real

continuum that was life Lamarcks separation of the establishment of relationships and

groupings in a system and identification of the plants in that system have been widely

accepted (but cf eg Godfray amp Knapp 2004 many of the contributors to Heywood et

al 2007) Perhaps the best way of identifying plants at the family level is by well-made

multiple access keys as in Watson and Dallwitz (1992a onwards family limits there may

differ substantially from those adopted here) Multiple access keys free users from the

constraints of dichotomous keys in which particular characters are needed at each step

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

20

of the identification process before there can be further progress Instead those

characters that happen to be evident on a specimen can be used in whatever order is

convenient when linked to illustrations glossaries etc their power is enormous (see

Dallwitz et al 2000 [2006] for the principles underlying their construction and use)

Nevertheless dichotomous keys such as those of Hutchinson (1973) and Franz Thonner

(Geesink et al 1981) have their uses Of course taxa coming out adjacent in keys may

well not be at all related

(When identifying large numbers of plants even more efficient than either style of

identification and certainly lots more fun is sight identification Unless you have a

photographic memory you have to build up your knowledge of comparative plant

morphology - on which the ability to make accurate identifications depends - by

repeated observation When faced with an unknown plant I always look for leaf teeth

and stipules distinctive hair types especially stellate and T-shaped hairs glands and

punctations of any sort the presence of latex or other exudate and check leaf insertion

smelling crushed leaves can also be helpful In this context nodal anatomy can usually

be checked using a razor and a hand lens or even simply carefully examining leaf scars

The short paragraphs added after most families may help in confirming familial

identifications)

For the record and for the little that it is worth there are 4 orders and 13 families

of gymnosperms characterised on these pages and together they include some 82

genera and 947 species For angiosperms comparable figures are 56 orders 445

families 13208 genera and 261750 species (of which monocots include 11 orders 89

families 2759 genera and 52760 species) Note however that higher mathematics

was never my strong point and anyway these are pretty meaningless figures even for

species which many (but not all) might concede smacked slightly more of reality than

other taxa estimates range as high as 422000 (Govaerts 2001) Furthermore numbers

of genera and species change daily Nevertheless as emphasized here families are

useful in teaching we as a community can ensure that their limits remain largely stable

and by concentrating on relatively few of them one can gain some familiarity with much

of the worlds flora For summaries of the sizes of orders and families and of the general

arrangement followed in these pages see also the Statistics page itself

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

21

A PHYLOGENETIC TREE OF SEED PLANTS (THE MAIN TREE)

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

22

The Main Tree above is a conservative summary of well supported relationships

between the major clades of extant seed plants Reading - interpreting and

understanding - such a phylogenetic tree is not necessarily easy so reading a tree may

require a bit of practice OHara (eg 1988 1992) provides an accessible introduction to

thinking and talking about history in the context of looking at phylogenetic trees More

recently Krell and Cranston (2004) Crisp and Cook (2005) and others have emphasized

how careful one must be when interpreting and talking about evolution in ladderized

trees in particular and phylogenetic trees in general Gregory (2008) provides a

particularly detailed discussion of this problem In ladderized trees the smaller (in terms

of numbers of terminals) sister taxon is consistently shown on the same side at every

node thus the trees tend to be pectinate like the teeth of a comb however the

horizontal axis that is as a result so evident has no polarity or particular direction The

use of the adjective basal when discussing phylogenetic trees is especially dangerous

(see also Wojciechowski et al 2004 D Soltis et al 2005b) When I use the term and the

context is not otherwise clear I am referring to the pectinations at the base of a

ladderised tree Remember that when talking about sister taxa one can never be basal

to the other or older or younger although members of one can be more derived - in the

sense of having more apomorphies - than the other (but even then one has to be

careful) The words primitive and advanced and lower and higher should

practically never be used especially when talking about taxa Amborellaceae and

Pinaceae are sister to all other flowering plants and Pinales and are basal to the crown

groups of all other flowering plantsPinales respectively but that does not mean they

are thereby primitive Pinaceae in particular have numerous apomorphies The word

plesiomorphic is far less loaded than primitive and can be used to talk about

individual characters

Most trees here have been more or less ladderized as is the one above The

ladderization may be imperfect for example you can see that on the Main Tree

asterids with ten orders follow rosids and relatives which have seventeen although

this is in part because of the currently unresolved nature of relationships between the

main clades of core eudicots (this is likely to change soon) The Main Tree could have

been drawn with Amborella Acorus or a host of other taxa at the far right without

offending any relationships phylogenetic trees are like mobiles the only fixed points

being the nodes Despite the problems just mentioned pectination interpreted

carefully has its value and it is not an arbitrary process As one reads the terminals of a

pectinate tree from left to right adjacent terminals will be separated by apomorphies

that will cumulate in an appropriate fashion When thinking of a book or a herbarium

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

23

sequence (see Haston et al 2007) this is of some value Nymphaeales and

Austrobaileyales are here adjacent on the tree but they could be separated by

hundreds of families in the sequence merely by rotating only the node from which

Nymphaeales and their sister group (all other angiosperms minus about 75 species)

arise If adjacent in a book or herbarium or as in these summary pages then it is

relatively easy to relate their characters both apomorphic and plesiomorphic but if

separated by hundreds of pages or two floors in a large building then it is less easy to

get anything from the sequence (Since all orders in the pages of the main part of this

site are preceded by the apomorphies of all nodes immediately below them in the seed

plant phylogeny and because of the linkages that have been built in to the site this

problem is much less serious there there is no sequence) So the particular sequence of

all seed plants that is based on a phylogenetic classification but is to be used in herbaria

or any other place where taxa have to be arranged linearly as in this part of APweb

will be one that maximizes the number of taxa that are both successive branches of the

tree and placed successively in the sequence Since specimens are generally filed under

families the outline of a new family sequence for arranging herbaria and books can now

be suggested (Haston et al 2007) and such a standardized sequence will further help

teaching and learning about plants

ON THINKING ABOUT APOMORPHIES

Identifying apomorphies is important because understanding the evolution of

morphology in the broad sense - ie including anatomy chemistry etc - in the context

of diversification is one of our major goals For identifying apomorphies several

preconditions must be met One needs to have an accurate well supported phylogeny

one has to have examined the right taxa both from the point of view of morphology and

molecules one has to have coded the characters right (ie delimited states

appropriately) and one has to use an appropriate model of evolution when fitting the

variation to the tree (see eg Omland 1999 Stevens 2006b) If the distributions of

apomorphies below are compared with those in other studies (eg Turgeon et al 2001

Bremer et al 2001 Endress 2001 Albach et al 2001a Judd amp Olmstead 2004 D Soltis

et al 2005b Zhang et al 2006 Judd et al 2007) the reader may find a number of

differences Although I have integrated such studies as far as is possible there are five

reasons that link to the preconditions just mentioned why there may be differences

where features are placed on trees

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

24

1 I may not have found all the information about a particular character there may be

disagreement over its interpretation or I have added information

2 The sampling of nearly all molecular studies is very incomplete (see Salisbury amp Kim

2001 for problems caused by sampling) indeed it is commonplace to decry the

incompleteness of molecular sampling But not only is the sampling in molecular studies

often less than we might wish that of the morphological and chemical characters whose

evolution we are interested in understanding is also often very poor So for many

anatomical chemical and embryological characters that are confidently said to

characterise families and other groups we all too often have no idea if those characters

are applicable to the whole clade or just to a subgroup within that clade However as

our sampling improves we can locate changes on the tree more precisely if in

unexpected positions Thus is is sometimes suggested that Ericaceae have ellagic acid

(Soltis et al 2005b) in fact the little evidence we have indicates that only a small clade

of some 80 species has ellagic acid and the rest of the family - itself alone about one

third of all Ericales - does not Similarly Ericaceae are noted for their

ectendomycorrhizal associations but Enkianthus sister to all other Ericaceae

apparently lacks these (Abe 2005) Thus these mycorrhizae and their associated

distinctive hair roots are unlikely to be apomorphies of Ericaceae - however it appears

that other families near Ericaceae may also have ectendomycorrhizae (see Asai 1934)

3 I am fitting characters to a very conservative tree with many polytomies although the

nodes that are utilised are for the most part strongly supported polytomies make the

optimisation of characters that is the assigment of character state change to a

particular node on the tree notably difficult (eg Madison amp Madison 2002) In nearly

all studies of the evolution of characters distributions of characters are optimised on a

more or less fully resolved tree and the construction of supertrees may yield yet more

detailed hypotheses of relationships (for literature on supertrees see Cotton amp

Wilkinson 2007 2008) Of course some nodes on such fully resolved trees andor

supertrees may have little support and optimisations of characters on such trees may

carry correspondingly little conviction Even parts of some of the trees used here have

poor support eg relationships within aquatic Alismatales etc although I indicate

when this is the case

4 Exactly how one goes about optimising a character on a tree is critically important

Even using simple parsimony optimisations (ACCTRAN or DELTRAN ACCelerated

TRANsitions or DELayed TRANsitions) the position of synapomorphies on trees - and

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

25

hence our ideas of evolution - may differ and this is still more true if one uses maximum

likelihood or Bayesian methods and makes apparently reasonable suggestions about

weighting gains over losses (eg Donoghue amp Ackerley 1996 Cunningham et al 1998

Omland 1997 1999 Ree amp Donoghue 1999 Polly 2001 Webster amp Purvis 2001

Ronquist 2004 Crisp amp Cook 2005) Sannier et al (2007) show how in in Arecaceae that

where on a phylogenetic tree one might peg changes in microsporogenesis will depend

on the methods one uses to do this and Pedersen et al (2007) discuss the sometimes

very substantial effect of node support on the posterior probabilities of ancestral

character states Here I use parsimony optimization not always as explicit as it might be

but I have often indicated where there are particularly important uncertainties as to the

positions of particular character changes on the tree

5 Finally although I have paid quite a lot of attention to the delimitation of the

character states that make up all the characterizations I have not spent enough time on

this critical operation If we are interested in understanding evolution then fitting the

basic variation - not character states - to a tree in principle allows greater flexibility in

understanding morphology in the context of local phylogenies (see also Stevens 2000

Endress 2005c) However many character states used here are delimited globally that

is they are circumscribed in the context of the variation shown by individual characters

across all angiosperms andor in the context of classic ideas of character evolution

Character states often have arbitrary limits and serve best to communicate

information whether they are in fact suitable for either phylogenetic analysis or

understanding evolution are separate issues Studies have rather unsurprisingly

perhaps but importantly shown that dividing the one character into different sets of

states may yield differing ideas of evolution of that character (eg Lamb Frye amp Kron

2003 Hibbett 2004) When looking at trees on which character states are optimised

one should bear in mind the problems surrounding the delimitation of states (eg

Stevens 2000 2006b) and the danger of using pollen or other types - constructs

based on many characters that vary independently but which effectively get lost in

these types thus Blackmore et al (2009) decomposed the pollen types in Asteraceae

into 52 characters

I use Remanes three main criteria of homology or better similarity when

determining the basic similarity of structures on different organisms (see Remane 1952)

These criteria are special properties position and intermediates Special properties

include anatomical or chemical characters gene expression data etc Position refers

to the position of an organ with respect to landmarks on the plant Although plants are

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

26

plants and landmarks are not as fixed as one might like trying to work out the relative

positions of parts is a good way of understanding morphology so long as one does not

try to out-do Procrustes Intermediates include intermediates found on the same

plant or on different plants Intermediates may observed during development or be

adult structures Thus one may find changes during development which suggest what

the nature of a particular structure is and very different stuctures on different plants

may turn out to be similar early in development Or one can simply compare different

taxa and see that structures that appear to be very different when comparing two

different taxa can be linked morphologically by looking at other taxa

Nevertheless the use of these criteria may not yield an unambiguous answer as to

what a structure is even given a solid phylogeny and an improved understanding of

development (see Jaramillo amp Kramer 2007 for a useful discussion) As Endress (2005c)

observed a number of features - position function development shape anatomy

histology gene activity and relationships to other taxa that clearly have petals - can be

used to distinguish a petal (for example) from other floral structures if a petal does not

have one of these features is it thereby not a petal Thus Maturen et al (2005) recently

found that floral organ diversity genes (B and C) were expressed in the large white

inflorescence bracts of Cornus (see also Costa et al 2005) Peney et al (2005) noted that

not all monosulcate pollen grains in monocots have the same developmental pathway

and that as a result such pollen might not have the same ancestral state Reeves and

Olmstead (2003) suggested that the genetic mechanisms causing monosymmetry in

Lamiales and Solanales were different and Serna and Martin (2006) described similar

problems with the development of hairs in Arabidopsis when compared with that of

hairs in Antirrhinum and Solanaceae Indeed as one perhaps might expect delimitation

of states and characters does not necessarily become easier with increasing knowledge

of development etc Thus Buzgo et al (2004) Matthews and Endress (2005) and others

have shown how hard it can be to distinguish between eg prophylls and other floral

structures as their behaviour is studied during the course of the development of the

flower Similarly at what concentration is a particular secondary metabolite deemed to

be present (Waterman (2007)

The validity of the approach used here that of fitting morphological variation to a

largely molecular-based tree may be questioned However I think it rather unlikely that

well-supported molecular branches will be overturned by morphological data Indeed

analyses of morphological data alone do provide support for many of the clades evident

in molecule-only analyses and analyses with morphological and molecular data

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

27

together may lead to increased support for clades (eg Hufford 1992 Nandi et al 1998

[but adding morphological data reduces support for a number of critical clades too]

Doyle amp Endress 2000 however in none of these papers is the use of morphology

without ambiguity) It is unfortunately clear that the use of morphology alone may not

suggest problems in the phylogenetic placement of taxa that later turn out to have been

wrongly included (eg Zhang et al 1992) Thus I have been wary of putting much weight

on clades that have only morphological support but note that I have treated molecule-

based clades with low bootstrap or jacknife support values (esp below 70) or low

posterior probabilies (below 095) likewise Although I may have been mistaken in

placing so much emphasis on molecular data in terms of providing the basic

phylogenetic framework for angiosperms morphological and molecular data are only

very rarely in irreconcilably strong conflict There are indeed a few places where the

conflict seems extreme These include the relative positions of the Monimiaceae and

Hernandiaceae (Laurales) the position of Hanguanaceae (Commelinales [as here] or

Zingiberales) and of Triplostegia (is it in Dipsacaceae or Valerianaceae - see

Dipsacales) Fossils are unlikely to affect the topologies of the trees presented here but

see below for their importance in understanding morphological evolution in general and

the evolution of angiosperms in particular Nevertheless some largely reject the idea

that trees based on molecular data alone can recover phylogenetic relationships

especially when branching points are old and prefer to used trees based on analysis of

morphological data including those taken from fossils (Hilton amp Bateman 2006 Farjon

2007)

All in all however the extent of the congruence between morphological and

molecular data is impressive and heartening and many clades can be characterised

morphologically It seemed in 1998 that there were no unambiguous morphological

synapomorphies for angiosperm orders (K Bremer 2000) and this is still true if by

unambiguous is meant non-homoplasious However many orders and other clades

have synapomorphies even if these may be indistinguishable at least at the current

level of morphological and developmental knowledge from parallel occurences

elsewhere As our morphological knowledge increases so too does the number of

apomorphies

Endress and Matthews (2006a) emphasize the importance of tendencies and

developmental constraints when thinking about characters of clades within the rosids -

many characters or character combinations occur in a rather sporadic fashion within

the clade and apparently notably less frequently outside it There are a number of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

28

examples of tendencies here and in the individual order pages such as the features

enclosed by parentheses in groups above the level of family eg see (cuticular waxes

as aggregated rodlets) for the commelinids and the discussion of the distribution of

polyandry that is flowers with numerous stamens in the asterid I + II groups etc

Tendencies may involve more than single characters When thinking of the

associations of features like integument thickness and vascularization of the integument

(Endress amp Matthews 2006a) one can imagine fairly simple developmental

preconditions being operative As Endress and Matthews (2006a) note it is difficult to

imagine vascular tissue developing in an integument only two cells thick Changes here

would be loosely correlated if morphologically linked However the strongly correlated

changes noted by Givnish et al (2005) are ecologically linked but are presumably

morphologicallydevelopmentally independent When there seem to be characters

evolving more or less together tests can be carried out to see if the changes are

concentrated on certain branches of the tree (eg Maddison 1990 Sanderson 1991

Maddison amp Maddison 2000) Such tests have rarely been carried out at higher levels in

angiosperms

One of the most striking examples of a tendency is the distribution of N-fixation

restricted as it is to a monophyletic group of four clades although it has arisen seven (or

perhaps several more) times independently within the N-fixing clade and several

members of two quite different kinds of bacteria are involved (eg Clawson et al 2004

Elliott et al 2007 Sprent amp James 2007) Other examples of work that bears on the issue

of tendencies include the findings that flowers of polysymmetrical Arabidopsis have

genes like TCP1 that are expressed asymmetrically during early development and TCP1

is a probable orthologue of the well-known CYC gene of Antirrhinum that is involved in

the development of monosymmetric slowers there (Cubas et al 2001 Costa et al 2005

etc) Parallelism might build on this underlying morphologically cryptic monosymmetry

even if details of the genetic mechanisms causing the monosymmetry evident in

particular groups may be different (Reeves amp Olmstead 2003 Cubas 2004 see above)

Similarly there have been several recent suggestions that the capability to synthesise a

particular metabolite may be switched off but not lost and so can sometimes be

reacquired (eg Wink amp Witte 1983 Wink 2003 Liscombe et al 2005 Larsson 2007

Waterman 2007) Hence perhaps the rather spotty distribution of many secondary

metabolites like ellagic acid the indole alkaloid camptothecin iridoids etc when

considered in the context of phylogenies As a non-botanical example - but a rather nice

one - Salwini-Plawen and Mayr (1961) suggested some time ago that there has been

considerable parallelism (40-65 or more independent origins) in the evolution of eyes in

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

29

metazoans However Pax 6 seems to be a master control gene that is involved in eye

formation perhaps throughout all bilateralians (eg Gehring amp Ikeo 1999 Erwin amp

Davidson 2002) which changes the nature of the problem Note however that in

general our invocation of evolutionary tendencies to explain the patterns of

distributions of characters is really an admission that we do not understand those

patterns

As we find out more about variation we will probably find fewer and fewer features

constant throughout a group Most unqualified statements of presence and absence

should properly be qualified as usually present or usually absent if one is thinking of

the characterisations on the order pages as encompassing the total variation within a

clade Thus Pistia alone among monocots as so far known has sieve tube plastids with

starch grains not protein crystals However this does not affect the fact that sieve tube

plastids with cuneate protein crystals and lacking starch grains are an apomorphy for

monocots Furthermore fossils in a number of cases suggest character combinations

unknown in extant taxa as may be seen in the discussions of Fagaceae Platanaceae

Iteaceae Calycanthaceae etc Confusing the issue there may be questions as to where

exactly on the tree a particular fossil is to be placed (eg see Nymphaeaceae

Calycanthaceae Archaefructus etc)

To summarize given our current understandings of both phylogenies and

characters evolution of some characters in which we are interested seems very labile

(see eg D Soltis et al 2005b Endress and Matthews 2006a Stevens 2006b Ekman et

al 2008 for a good recent study) and I have been cautious when talking about character

evolution Much effort must continue to be spent in summarizing characters of well-

established clades at all levels providing features by which they may be recognized and

signaling synapomorphies Remember that (1) the basic morphological anatomical and

chemical knowledge of many critical taxa is woefully incomplete (2) different

assumptions about character evolution may greatly affect the position of

synapomorphies on trees (3) in many cases relationships within and between many

groups are too uncertain at present to worry very much about synapomorphies and

(4) we must be clear about what we do and do not not know As mentioned above I

have indicated in a number of places where there are particularly important

uncertainties as to where characters should be placed on the tree Nevertheless it is a

relatively easy matter to update notes such as these and it can be a simple matter to

incorporate new data on characters that have never before been considered in the

context of a tree Much basic - and unfortunately perhaps unfashionable - work must be

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

30

carried to clarify the distribution of morphological anatomical and chemical characters

for examples of what can to be done see the work of P K Endress and collaborators

(floral morphology and development) and S R Jensen and collaborators (iridoids) But

acquiring information about nodal anatomy cell and tissue distribution nodal anatomy

and the like is not difficult at all and should be generally encouraged

But all this is merely a necesary prelude to the understanding of evolution For this

one needs to know a lot more including dating the phylogeny understanding the

function(s) of characters factoring in relevant aspects of the palaeoenvironment in

which an apomorphy first appeared etc

SUMMARY OF THE SYSTEM

Below is a formalised summary of the relationships within orders of the families of seed

plants There are a few families that are not recognised even as options in APG II and

vice versa as well as a few extra orders The families to be recognised in the most

recent edition of Mabberleys The Plant Book (Mabberley 2008) are also largely

consistent with those below But all differences are trivial and will - I hope - eventually

disappear

Square brackets - [] - enclose clades the plus sign - + - designates sister taxa a

comma - - denotes part of a polytomy and quotation marks - - denotes a

paraphyletic group

SEED PLANTS

GYMNOSPERMS

Cycadales

Cycadaceae + Zamiaceae

Ginkgoales

Ginkgoaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

31

Pinales

Pinaceae [[Araucariaceae [Podocarpaceae + Phyllocladaceae]] [Sciadopityaceae

[Taxaceae + Cupressaceae]]]

Gnetales

Ephedraceae [Gnetaceae + Welwitschiaceae]

ANGIOSPERMSFLOWERING PLANTS

Amborellales

Amborellaceae

Nymphaeales

Hydatellaceae [Cabombaceae + Nymphaeaceae]

Austrobaileyales

Austrobaileyaceae [Schisandraceae + Trimeniaceae]

Chloranthales

Chloranthaceae

MAGNOLIIDS

Magnoliales

Myristicaceae [Magnoliaceae [[Himantandraceae + Degeneriaceae] [Eupomatiaceae +

Annonaceae]]]

Laurales

Calycanthaceae [[Siparunaceae [Gomortegaceae + Atherospermataceae]] [Monimiaceae

[Hernandiaceae + Lauraceae]]]

Canellales

Canellaceae + Winteraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

32

Piperales

[Hydnoraceae + Aristolochiaceae] [Piperaceae + Saururaceae]

MONOCOTS

Acorales

Acoraceae

Alismatales

Araceae [Tofieldiaceae [[Alismataceae [Hydrocharitaceae + Butomaceae]]

[Scheuchzeriaceae [Aponogetonaceae [Juncaginaceae [Maundiaceae [[Posidoniaceae

[Ruppiaceae + Cymodoceaceae]] [Zosteraceae + Potamogetonaceae]]]]]]]

Petrosaviales

Petrosaviaceae

Dioscoreales

Nartheciaceae [[Taccaceae + Thismiaceae] [Burmanniaceae + Dioscoreaceae]]

Pandanales

Velloziaceae Triuridaceae Stemonaceae [Pandanaceae + Cyclanthaceae]

Liliales

Corsiaceae [Campynemataceae [Petermanniaceae [Colchicaceae + Alstroemeriaceae]]

Melanthiaceae [[Philesiaceae + Rhipogonaceae] [Smilacaceae + Liliaceae]]]

Asparagales

Orchidaceae [[Boryaceae [Blandfordiaceae [Lanariaceae [Asteliaceae + Hypoxidaceae]]]]

[[Ixioliriaceae + Tecophilaeaceae] [Doryanthaceae [Iridaceae [Xeronemataceae

[Xanthorrhoeaceae [Alliaceae + Asparagaceae]]]]]]]

COMMELINIDS

Unplaced

Dasypogonaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

33

Arecales

Arecaceae

Poales

[Typhaceae + Bromeliaceae] [Rapateaceae [[Thurniaceae [Juncaceae + Cyperaceae]]

[[Mayacaceae [Eriocaulaceae + Xyridaceae]] [Flagellariaceae [Anarthriaceae

[Centrolepidaceae + Restionaceae]]] [Joinvilleaceae [Ecdeiocoleaceae + Poaceae]]]]]]]

Commelinales

[Commelinaceae + Hanguanaceae] [Philydraceae [Haemodoraceae + Pontederiaceae]]

Zingiberales

Musaceae [Strelitziaceae + Lowiaceae] Heliconiaceae [[Cannaceae + Marantaceae]

[Costaceae + Zingiberaceae]]

Ceratophyllales

Ceratophyllaceae

EUDICOTS

Ranunculales

Eupteleaceae [Papaveraceae [[[Lardizabalaceae + Circaeasteraceae] [Menispermaceae

[Berberidaceae + Ranunculaceae]]]]

Sabiales

Sabiaceae

Proteales

Nelumbonaceae [Platanaceae + Proteaceae]

Trochodendrales

Trochodendraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

34

Buxales

Haptanthaceae [Buxaceae + Didymelaceae]

CORE EUDICOTS

Gunnerales

Gunneraceae + Myrothamnaceae

Dilleniales

Dilleniaceae

Saxifragales

Peridiscaceae [[Paeoniaceae [Altingiaceae [Hamamelidaceae [Cercidiphyllaceae +

Daphniphyllaceae]]]] [[Crassulaceae [Aphanopetalaceae [Tetracarpaeaceae

[Penthoraceae + Haloragaceae]]]] [Iteaceae [Grossulariaceae + Saxifragaceae]]]]

Cynomoriaceae unplaced

Vitales

Vitaceae

ROSIDS

FABIDROSID I

Zygophyllales

Krameriaceae + Zygophyllaceae

Celastrales

Lepidobotryaceae + Celastraceae

Oxalidales

Huaceae [[Connaraceae + Oxalidaceae] [Cunoniaceae [Elaeocarpaceae [Brunelliaceae +

Cephalotaceae]]]]

Malpighiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

35

[Achariaceae [Goupiaceae [Violaceae + Passifloraceae] [Lacistemataceae Salicaceae]]

[Lophopyxidaceae Putranjivaceae] [Ctenolophonaceae [Erythroxylaceae +

Rhizophoraceae]] Linaceae Ixonanthaceae Humiriaceae Irvingiaceae

Centroplacaceae Pandaceae [[Bonnetiaceae + Clusiaceae] [Calophyllaceae

[Hypericaceae + Podostemaceae]]] [Malpighiaceae + Elatinaceae] Ochnaceae

[Peraceae [Rafflesiaceae + Euphorbiaceae]] [Phyllanthaceae + Picrodendraceae]

[Balanopaceae [[Trigoniaceae + Dichapetalaceae] [Chrysobalanaceae +

Euphroniaceae]]] Caryocaraceae

N-FIXING CLADE

Fabales

Quillajaceae [Fabaceae [Polygalaceae + Surianaceae]]

Rosales

Rosaceae [[Barbeyaceae [Elaeagnaceae Dirachmaceae Rhamnaceae]] [Ulmaceae

[Cannabaceae [Moraceae + Urticaceae]]]]

Cucurbitales

Anisophylleaceae [[Corynocarpaceae + Coriariaceae] [Cucurbitaceae [Tetramelaceae

[Datiscaceae + Begoniaceae]]]] Apodanthaceae

Fagales

Nothofagaceae [Fagaceae [[Myricaceae + Juglandaceae] [Casuarinaceae

[Ticodendraceae + Betulaceae]]]]

MALVIDROSID II

pgtGeraniales

Geraniaceae [[Melianthaceae + Francoaceae] [Vivianaceae + Ledocarpaceae]]

Myrtales

Combretaceae [[Onagraceae + Lythraceae] [[Vochysiaceae + Myrtaceae]

[Melastomataceae [Crypteroniaceae [Alzateaceae + Penaeaceae]]]]]

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

36

Crossosomatales

[Staphyleaceae [Guamatelaceae [Crossosomataceae + Stachyuraceae]]] [Aphloiaceae

[Geissolomataceae + Strasburgeriaceae]]

Picramniales

Picramniaceae

Sapindales

Biebersteiniaceae Nitrariaceae [Kirkiaceae [Anacardiaceae + Burseraceae]]

[Sapindaceae [Simaroubaceae Rutaceae Meliaceae]]

Huerteales

Gerradinaceae [Dipentodontaceae + Tapisciaceae]

Malvales

Neuradaceae [Thymelaeaceae [Sphaerosepalaceae Bixaceae [Cistaceae

[Sarcolaenaceae + Dipterocarpaceae]] [Cytinaceae + Muntingiaceae] Malvaceae]]

Brassicales

[Akaniaceae + Tropaeolaceae] [[Moringaceae + Caricaceae] [Setchellanthaceae

[Limnanthaceae [[Koeberliniaceae [Bataceae + Salvadoraceae]] [Emblingiaceae

[Pentadiplandraceae [Gyrostemonaceae Resedaceae Stixis and relatives] Tovariaceae

[Capparaceae [Cleomaceae + Brassicaceae]]]]]]]]

Berberidopsidales

Aextoxicaceae + Berberidopsidaceae

Santalales

Erythropalaceae Olacaceae [[Loranthaceae [Misodendraceae + Schoepfiaceae]]

[Opiliaceae + Santalaceae]] Balanophoraceae unplaced

Caryophyllales

[[Droseraceae [Nepenthaceae [Drosophyllaceae [Ancistrocladaceae +

Dioncophylleaceae]]]]] [[Frankeniaceae + Tamaricaceae] [Polygonaceae +

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

37

Plumbaginaceae]]] [Rhabdodendraceae [Simmondsiaceae [[Asteropeiaceae +

Physenaceae] [[Caryophyllaceae [Achatocarpaceae + Amaranthaceae]]

[Stegnospermataceae [Limeaceae [[Lophocarpaceae [Barbeuiaceae [Aizoaceae

[Sarcobataceae Phytolaccaceae Nyctaginaceae]]]] [Molluginaceae [Halophytaceae

Montiaceae Basellaceae Didiereaceae [Talinaceae [Portulacaceae [Anacampseros etc +

Cactaceae]]]]]]]]]]]]]

ASTERIDS

Cornales

[Cornaceae + Nyssaceae] [Hydrangeaceae + Loasaceae] Hydrostachyaceae [Curtisiaceae

+ Grubbiaceae]

Ericales

[Balsaminaceae [Marcgraviaceae + Tetrameristaceae]] [[Polemoniaceae +

Fouquieraceae] Lecythidaceae [[Sladeniaceae + Pentaphylacaceae] [Sapotaceae

[Ebenaceae [Maesaceae [Theophrastaceae [Myrsinaceae + Primulaceae]]]]]

[Mitrastemonaceae Theaceae [Symplocaceae [Styracaceae + Diapensiaceae]]

[[Sarraceniaceae [Roridulaceae + Actinidiaceae] [Clethraceae [Cyrillaceae +

Ericaceae]]]]]]

LAMIIDASTERID I

Unplaced

Metteniusaceae Oncothecaceae Icacinaceae and some unplaced ex-Icacinaceae

Garryales

Garryaceae + Eucommiaceae

Unplaced

Boraginaceae Vahliaceae

Gentianales

Rubiaceae [Gentianaceae [Loganiaceae [Gelsemiaceae + Apocynaceae]]]

Lamiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

38

Plocospermataceae [Oleaceae [Tetrachondraceae [Calceolariaceae

[Gesneriaceae Peltanthera [Pedaliaceae Carlemanniaceae Martyniaceae Stilbaceae

Plantaginaceae Linderniaceae Bignoniaceae Verbenaceae Lamiaceae Paulowniaceae

Schlegeliaceae Thomandersiaceae Phrymaceae Rehmannia Acanthaceae

Scrophulariaceae Orobanchaceae Byblidaceae Lentibulariaceae]]]]]

Solanales

[Montiniaceae [Sphenocleaceae + Hydroleaceae]] [Convolvulaceae + Solanaceae]

CAMPANULIDASTERID II

Aquifoliales

[[Cardiopteridaceae + Stemonuraceae] [Aquifoliaceae [Helwingiaceae +

Phyllonomaceae]]

Asterales

[Rousseaceae + Campanulaceae] Pentaphragmataceae [[Alseuosmiaceae [Phellinaceae

+ Argophyllaceae]] Stylidiaceae [Menyanthaceae [Goodeniaceae [Calyceraceae +

Asteraceae]]]]

Escalloniales

Escalloniaceae

Bruniales

[Bruniaceae + Columelliaceae]

Apiales

Pennantiaceae [Torricelliaceae [Griseliniaceae [Pittosporaceae [Araliaceae

[Myodocarpaceae + Apiaceae]]]]]

Paracryphiales

Paracryphiaceae

Dipsacales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

39

Adoxaceae [Diervillaceae [Caprifoliaceae [Linnaeaceae [Morinaceae [Dipsacaceae +

Valerianaceae]]]]]

LINKS TO ORDERS AND FAMILIES

[Back to Top]

MAIN GROUPINGS

Asterids commelinids core eudicots asterid 1 asterid

2 eudicots gymnosperms Magnoliophyta monocots N-fixing claderosids seed plants

ALPHABETICAL LISTING OF ALL ORDINAL NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | L | M | N | O | P | Q | R | S | T | U | V | W | X | Z

Abietales Acanthales Acerales Acorales Actinidiales Actinostrobales Adoxales Aescul

ales Agavales Aizoales AkanialesAlismatales Alliales Alseuosmiales Alstroemeriales

Altingiales Amaranthales Amaryllidales Amborellales AmbrosialesAmmiales Amomal

es Ancistrocladales Anisophylleales Annonales Anthobolales Apiales Apocynales Apo

nogetonalesAquifoliales Arales Araliales Aralidiales Araucariales Arecales Aristolochi

ales Athrotaxidales Asarales AsclepiadalesAsparagales Asphodelales Asparagales Ast

eliales Atriplicales Aucubales Austrobaileyales Avenales

Balanitales Balanopales Balanophorales Balsaminales Barbeyales Barclayales Batales

Begoniales BerberidalesBerberidopsidales Betulales Biebersteiniales Bignoniales Bix

ales Boraginales Brassicales Brexiales Bromeliales BrunialesBrunoniales Burmannial

es Burserales Butomales Buxales Byblidales

Cactales Callitrichales Calycanthales Calycerales Campanulales Campynematales Can

ellales Cannales CapparalesCaprifoliales Cardiopteridales Carduales Caricales Carle

manniales Caryophyllales Cassiales Casuarinales CelastralesCentrolepidales Cephalot

ales Cephalotaxales Ceratophyllales Cercidiphyllales Chenopodiales Chironiales Chlor

anthalesChrysobalanales Cinchonales Circaeasterales Cistales Citrales Cocosales Col

chicales Columelliales CombretalesCommelinales Connarales Convolvulales Coriarial

es Cornales Corylales Corynocarpales Crassulales CrossosomatalesCucurbitales Cunn

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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inghamiales Cunoniales Cupressales Cycadales Cyclanthales Cymodoceales Cynarales

CynomorialesCyperales Cyrillales Cytinales

Daphnales Daphniphyllales Dasypogonales Datiscales Degeneriales Desfontainiales D

iapensiales Didymelales DillenialesDioales Dioncophyllales Dioscoreales Diospyrales

Dipentodontales Dipsacales Droserales

Ebenales Echiales Elaeagnales Elaeocarpales Elatinales Elodeales Emmotales Empetr

ales Ephedrales Ericales EriocaulalesErythropalaless Escalloniales Eucommiales Euph

orbiales Eupomatiales Eupteleales Euryalaless

Fabales Fagales Falcatifoliales Ficales Flacourtiales Flagellariales Fouquieriales Franc

oales Frangulales

Galiales Garryales Geissolomatales Gentianales Geraniales Gesneriales Ginkgoales G

laucidiales Globulariales GnetalesGoodeniales Greyiales Griseliniales Grossulariales

Grubbiales Gunnerales Gyrocarpales Gyrostemonales

Haemodorales Haloragales Hamamelidales Hanguanales Heisteriales Helleborales He

lwingiales Himantandrales HippuridalesHomaliales Hortensiales Huales Huerteales

Hydatellales Hydnorales Hydrangeales Hydrastidales HydrocharitalesHydropeltidales

Hydrostachyales Hypericales Hypoxidales

Icacinales Illiciales Iridales Irvingales Iteales Ixiales

Jasminales Juglandales Julianiales Juncaginales Juncales

Lacistematales Lactoridales Lamiales Lardizabalales Laurales Lecythidales Ledocarpal

es Leitneriales LentibularialesLigustrales Liliales Limnanthales Linales Liriales Loasal

es Lobeliales Loganiales Lonicerales Loranthales LowialesLythrales

Magnoliales Malpighiales Malvales Marathrales Marcgraviales Mayacales Medusagy

nales Medusandrales MelanthialesMelastomatales Meliales Melianthales Meliosmal

es Menispermales Menyanthales Metteniusales MitrastemonalesMiyoshiales Moni

miales Moringales Musales Myricales Myristicales Myrothamnales Myrsinales Myrta

les

Najadales Nandinales Narcissales Nartheciales Nelumbonales Nepenthales Neuradal

es Nitrariales Nolanales NothofagalesNyctaginales Nymphaeales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Ochnales Oenotherales Olacales Oleales Onagrales Oncothecales Opuntiales Orchid

ales Oxalidales

Saxifragales Pandales Pandanales Papaverales Paracryphiales Parasitaxales Paridales

Parnassiales PassifloralesPenaeales Pennantiales Pentaphragmatales Peridiscales P

etiveriales Petrosaviales Phellinales Philydrales PhyllanthalesPhysenales Phytolaccal

es Picramniales Pinales Pinguiculales Piperales Pittosporales Plantaginales Platanale

sPlumbaginales Poales Podocarpales Podophyllales Podostemales Polemoniales Pol

ygalales Polygonales PontederialesPortulacales Posidoniales Potamogetonales Primu

lales Proteales

Quercales Quillajales Quintiniales

Rafflesiales Ranunculales Rapateales Resedales Restionales Rhabdodendrales Rhamn

ales Rhinanthales RhizophoralesRhodorales Rhoipteleales Roridulales Rosales Rouss

eales Rubiales Ruppiales Rutales

Sabiales Salicales Salvadorales Samolales Samydales Sanguisorbales Santalales Sapin

dales Sapotales SarracenialesSaxegotheales Saxifragales Scheuchzeriales Sciadopitya

les Scleranthales Scrophulariales Scyphostegiales SedalesSilenales Simmondsiales S

milacales Solanales Sphenocleales Sphenostemonales Stangeriales Stellariales Stemo

nalesStilbales Stylidiales Styracales Surianales

Taccales Tamales Tamaricales Taxales Taxodiales Tecophilaeales Terebinthales Tern

stroemiales Theales TheligonalesThymelaeales Tiliales Tofieldiales Torricelliales Tov

ariales Tribelales Trilliales Trimeniales Triuridales TrochodendralesTropaeolales Turn

erales Typhales

Ulmales Urticales

Vacciniales Vahliales Vallisneriales Velloziales Veratrales Verbenales Viburnales Vinc

ales Violales Viscales VitalesVochysiales

Welwitschiales Winterales

Xanthorrhoeales Ximeniales Xyridales

Zamiales Zingiberales Zosterales Zygophyllales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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ALPHABETICAL LISTING OF ALL FAMILY NAMES OF SEED PLANTS WITH LINKS

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which the genus of the basionym is to be found

Abietaceae Abolbodaceae Abrophyllaceae Acalyphaceae Acanthaceae [Acanthochla

mydaceae] Acarnaceaee AceraceaeAchariaceae Achatocarpaceae Achradaceae Achy

ranthaceae Acmopylaceae Acoraceae Acristaceae ActaeaceaeActinidiaceae Actinota

ceae Amentotaxaceae Adenogrammaceae Adoxaceae Aegialitidaceae Aegicerataceae

Aegilopaceae Aeginetiaceaee Aegiphilaceae Aesculaceae Aextoxicaceae Agapanthac

eae Agavaceae AgdestidaceaeAgialidaceae Agrimoniaceae Agrostidaceae Ailanthace

ae Aitoniaceae Aizoaceae Akaniaceae Alangiaceae AlchemillaceaeAldrovandaceae A

lismataceae Alliaceae Allioniaceae Allophylaceae Aloaceae Alopecuraceae Alpiniacea

e AlseuosmiaceaeAlsinaceae Alsodeiaceae Alstroemeriaceae Altingiaceae Alzateacea

e Amaranthaceae Amaryllidaceae Amborellaceae Ambrosiaceae Amentotaxaceae A

mmanniaceae Ammiaceae Amomaceae Ampelidopsaceae Amygdalaceae Amyridacea

eAnacardiaceae Anagallidaceae Anarthriaceae Anchusaceae Ancistrocladaceae Andr

omedaceae AndropogonaceaeAndrostachyaceae Androsynaceae Anemarrhenaceae

Anemonaceae Angelicaceae Anisophylleaceae AnnonaceaeAnomochloaceae Anopter

aceae Anrederaceae Anthemidaceae Anthericaceae Anthobolaceae Antidesmataceae

Antirrhinaceae Antoniaceae Aparinaceae Aphanopetalaceae Aphloiaceae Aphyllanth

aceae Apiaceae ApocynaceaeApodanthaceae Aponogetonaceae Aporusaceae Apose

ridaceae Apostasiaceae Aptandraceae AquifoliaceaeAquilariaceae Aquilegiaceae A

raceae Aragoaceae Araliaceae Aralidiaceaee Araucariaceae Arbutaceae Arceuthidace

aeArceuthobiaceae Arctostaphylaceae Arctotidaceae Ardisiaceae Arecaceae Argoph

yllaceae Arisaraceae AristolochiaceaeAristoteliaceae Arjonaceae Armeriaceae Artem

isiaceae Arthrotaxidaceae Artocarpaceae Arundinaceae ArundinellaceaeAsaraceae A

sclepiadaceae Ascyraceaee Asiraceae Asparagaceae Asperulaceae Asphodelaceae As

pidistraceaeeAsteliaceae Asteraceae Asteranthaceae Asteropeiaceae Astragalaceae

Astrocarpaceae AthanasiaceaeAtherospermataceae Atriplicaceae Atropaceae Aucuba

ceae Aurantiaceae Austrobaileyaceae Austrotaxaceae AvenaceaeAverrhoaceae Avet

raceae Avicenniaceae Azaleaceae Azimaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Balanitaceae Balanopaceae Balanophoraceae Balsameaceae Balsaminaceae Bambusa

ceae Barbaceniaceae BarbeuiaceaeBarbeyaceae Barclayaceae Barringtoniaceae Base

llaceae Bataceae Baueraceae Bauhiniaceae Baxteriaceae BegoniaceaeBehniaceae B

elangeraceae Belloniaceae Bembiciaceae Berberidaceae Berberidopsidaceae Berryac

eaee BersamaceaeBertyaceae Berzeliaceae Besleriaceae Betaceae Betulaceae Biebe

rsteiniaceae Bifariaceae Bignoniaceae BischofiaceaeBixaceae Blakeaceae Blandfordia

ceae Blattiaceae Blepharocaryaceae Blitaceae Blyxaceae Boerlagellaceae Bolivaracea

eBombacaceae Bonnetiaceae Bontiaceae Boopidaceae Boraginaceae Borassaceae B

oroniaceae BoryaceaeBotryodendraceae Bougainvilleaceae Boweniaceae Brassicacea

e Brachycaulaceae Bretschneideraceae BrexiaceaeBromeliaceae Brownlowiaceae Br

unelliaceae Bruniaceae Brunoniaceae Brunsvigiaceae Bryoniaceae BuchneraceaeBuci

daceae Buddlejaceae Buglossaceae Bulbocodiaceae Bumeliaceae Bupleuraceae Burc

hardiaceae BurmanniaceaeBurseraceae Butneriaceae Butomaceae Buxaceae Byblida

ceae Byttneriaceae

Cabombaceae Cacaoaceae Cactaceae Caesalpiniaceae Caladiaceae Calamaceae Calce

olariaceae CalectasiaceaeCalendulaceae Callaceae Callicomaceae Calligonaceae Calli

traceae Callitrichaceae Calochortaceae CalophyllaceaeCalthaceae Calycanthaceae Ca

lyceraceae Cambogiaceae Camelliaceae Campanulaceae CampynemataceaeCanacom

yricaceae Canellaceae Cannabaceae Cannaceae Canopodaceae Canotiaceae Cansjera

ceae CapparaceaeCaprariaceae Caprifoliaceae Cardiopteridaceae Carduaceae Carica

ceae Carissaceae Carlemanniaceae CarpinaceaeCarpodetaceae Cartonemataceae Ca

ryocaraceae Caryophyllaceae Casearieaceae Cassiaceae CassipoureaceaeCassythacea

e Castaneaceae Castelaceae Casuarinaceae Catesbaeaceae Cathedraceae Cecropiace

ae CedraceaeCedrelaceae Celastraceae Celosiaceae Celtidaceae Centaureaceae Cen

trolepidaceae Centroplacaceae CepaceaeCephalanthaceae Cephalotaceae Cephalota

xaceae Cerastiaceae Ceratoniaceae Ceratophyllaceae CerberaceaeCercidiphyllaceae

Cercocarpaceae Cercodiaceae Cereaceae Cerinthaceae Ceroxylaceae Cestraceae Cev

alliaceaeChailletiaceae Chamaedoraceae Chamaemoraceae Chamelauciaceae Chaun

ochitonaceae Chelidoniaceae ChelonaceaeChenopodiaceae Chimonanthaceae Chingi

thamnaceae Chionographidaceae Chiranthodendraceae ChironiaceaeChlaenaceae Ch

loanthaceae Chloranthaceae Chloridaceae Chlorogalaceae Chrysobalanaceae Cicerac

eae CichoriaceaeCimicifugaceae Cinchoniaceae Circaeaceae Circaeasteraceae Cissac

eae Cistaceae Citraceae Clematidaceae CleomaceaeClethraceae Clusiaceae Cneorac

eae Cnestidaceae Cnicaceae Cobaeaceae Cochlospermaceae Cocosaceae Coffeaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Colchicaceae Coleogynaceae Columelliaceae Combretaceae Commelinaceae Comocl

adiaceae Compositae CompsoaceaeConantheraceae Connaraceae Conostylidaceae C

onvallariaceae Convolvulaceae Cordiaceae CoreopsidaceaeCoriandraceae Coriariacea

e Coridaceae Corispermaceae Cornaceae Corokiaceae Coronillaceae Correaceae Cor

rigiolaceaeCorsiaceae Corydalaceae Corylaceae Corynocarpaceae Coryphaceae Cost

aceae Cotyledonaceae CoulaceaeCoutariaceae Coutoubiaceae Crassulaceae Crescen

tiaceae Cressaceae Crinaceae Crocaceae CroomiaceaeCrossosomataceae Crotonacea

e Cruciferae Cryptaceae Crypteroniaceae Cryptocorynaceae CryptomeriaceaeCtenol

ophonaceae Cucurbitaceae Cunninghamiaceae Cunoniaceae Cupressaceae Curcumac

eae Curtisiaceae CuscutaceaeCyananthaceae Cyanastraceae Cynanchaceae Cyanella

ceae Cycadaceae Cyclanthaceae CyclantheraceaeCyclocheilaceae Cydoniaceae Cymo

doceaceae Cynaraceae Cynocrambaceae Cynomoriaceae Cyperaceae CyphiaceaeCyp

hocarpaceae Cypripediaceae Cyrillaceae Cyrtandraceae Cyrtanthaceae Cytinaceae

Dacrycarpaceae Dactylanthaceae Damasoniaceae Daphnaceae Daphniphyllaceae Das

ypogonaceae DatiscaceaeDaturaceae Daucaceae Davidiaceae Davidsoniaceae Decais

neaceae Deeringiaceae Degeneriaceae DelphiniaceaeDendrophthoaceae Columelliac

eae Detariaceae Dialypetalanthaceae Dianellaceae Dianthaceae DiapensiaceaeDicha

petalaceae Dichondraceae Diclidantheraceae Dicrastylidiaceae Dictamnaceae Didiere

aceae DidymelaceaeDidymocarpaceae Diegodendraceae Diervillaceae Digitalidaceae

e Dilatridaceae Dilleniaceae Dioaceae DionaeaceaeDioncophyllaceae Dioscoreaceae

Diosmaceae Diospyraceae Dipentodontaceae Diphylleiaceae Diplolaenaceae Dipsaca

ceaeDipterocarpaceae Dirachmaceae Disanthaceae Diselmaceae Dodonaeaceae Do

mbeyaceae Donatiaceae DorsteniaceaeDortmannaceae Doryanthaceae Drabaceae D

racaenaceae Dracontiaceae Drimyidaceae Droseraceae DrosophyllaceaeDryadaceae

Duabangaceae Duckeodendraceae Dulongiaceae Durantaceae Durionaceae Dysphani

aceae

Ebenaceae Eccremidaceae Ecdeiocoleaceae Echiaceae Echinopaceae Ehretiaceae Ela

eagnaceae ElaeocarpaceaeElegiaceae Elatinaceae Ellisiaceae Ellisiophyllaceae Elodea

ceae Elytranthaceae Embeliaceae Emblingiaceae EmmotaceaeEmpetraceae Encepha

lartaceae Engelhardtiaceae Enhalaceae Epacridaceae Ephedraceae Ephemeranthacea

eEpilobiaceae Epimediaceae Eragrostidaceae Eremolepidaceae Eremosynaceae Erica

ceae Erinaceae EriocaulaceaeEriogonaceae Eriospermaceae Erodiaceae Erycibaceae

Eryngiaceae Erysimaceae Erythroniaceae ErythropalaceaeErythrospermaceae Erythro

xylaceae Escalloniaceae Eschscholziaceae Eucomidaceae Eucommiaceae Eucryphiace

aeEuonymaceae Eupatoriaceae Euphorbiaceae Euphrasiaceae Euphroniaceae Eupo

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matiaceae Eupteleaceae EuryalaceaeEustrephaceae Euthemidaceae Eutocaceae Exb

ucklandiaceae Exocarpaceae

Fabaceae Fagaceae Falcatifoliaceae Ferulaceae Festucaceae Ficaceae Ficoidaceae Fit

zroyaceae FlacourtiaceaeFlagellariaceae Flindersiaceae Foetidiaceae Forestieraceae

Fothergillaceae Fouquieriaceae Fragariaceae FrancoaceaeFrangulaceae Frankeniacea

e Fraxinaceae Fraxinellaceae Freycinetiaceae Fritillariaceae Fuchsiaceae Fumariacea

eFunkiaceae

Gaiadendraceae Galacaceae Galanthaceae Galaxiaceae Galedupaceae Galeniaceae G

aliaceae Garciniaceae GardeniaceaeGardneriaceae Garryaceae Geissolomataceae Ge

itonoplesiaceae Gelsemiaceae Geniostomaceae GentianaceaeGeonomataceae Geosir

idaceae Geraniaceae Gerrardinaceae Gesneriaceae Gethyllidaceae Gilliesiaceae Gina

lloaceaeGinkgoaceae Gisekiaceae Gladiolaceae Glaucidiaceae Glechomaceae Glinace

ae Globulariaceae Gnaphaliaceae GnetaceaeGoetzeaceae Gomortegaceae Gomphiac

eae Gomphrenaceae Gonystylaceae Goodeniaceae Gordoniaceae GouaniaceaeGoup

iaceae Gramineae Gratiolaceae Grewiaceae Greyiaceae Grielaceae Griseliniaceae Gr

onoviaceae GrossulariaceaeGrubbiaceae Guaiacanaceae Guamatelaceae Guettardac

eae Gunneraceae Gustaviaceae Guttiferae GyrocarpaceaeGyrostemonaceae

Hachetteaceae Haemanthaceae Haemodoraceae Halesiaceae Hallieraceae Halocarpa

ceae Halophilaceae HalophytaceaeHaloragaceae Hamamelidaceae Hanguanaceae H

aptanthaceae Harmandaceae Hebenstretiaceae HectorellaceaeHederaceae Hedyosm

aceae Hedyotidaceae Hedysaraceae Heisteriaceae Heleniaceae Heliamphoraceae H

elianthaceaeHelianthemaceae Helichrysaceae Heliconiaceae Helicteraceae Heliotropi

aceae Helleboraceae Heloniadaceae HelosidaceaeHelwingiaceae Hemerocallidaceae

Hemimeridaceae Hemiphylacaceae Henriqueziaceae Henslowiaceae Hermanniaceae

Hernandiaceae Herniariaceae Herreriaceae Hesperocallidaceae Heterantheraceae He

teropyxidaceae HeterostylaceaeHibbertiaceae Hibiscaceae Hilleriaceae Himantandra

ceae Hippocastanaceae Hippocrateaceae HippomanaceaeHippophaeaceae Hippurida

ceae Hirtellaceae Holacanthaceae Homaliaceae Hopkinsiaceae Hoplestigmataceae H

ordeaceaeHornschuchiaceae Hortensiaceae Hortoniaceae Hostaceae Houstoniaceae

Huaceae Huerteaceae HugoniaceaeHumbertiaceae Humiriaceae Hyacinthaceae Hyd

atellaceae Hydnoraceae Hydrangeaceae Hydrastidaceae HydrillaceaeHydrocharitacea

e Hydrocotylaceae Hydrogetonaceae Hydroleaceae Hydropeltidaceae Hydrophylacac

eae HydrophyllaceaeHydrostachyaceae Hymenocardiaceae Hypecoaceae Hyperanth

eraceae Hypericaceae Hypopityaceae HypoxidaceaeHypseocharitaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta

ceae ImperatoriaceaeInocarpaceae Inulaceae Ionidiaceae Iriarteaceae Iridaceae Irvi

ngiaceae Isnardiaceae Isophysidaceae IteaceaeStrasburgeriaceae Ixiaceae Ixioliriace

ae Ixonanthaceae

Jalapaceae Jamboliferaceae Japonoliriaceae Jasionaceae Jasminiaceae Johnsoniaceae

Joinvilleaceae JuglandaceaeJulianiaceae Juncaceae Juncaginaceae Juniperaceae Jus

siaeaceae Justiciaceae

Kadsuraceae Kaliphoraceae Kalmiaceae Kaniaceae Kiggelariaceae Kingdoniaceae Kin

giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae

Krameriaceae

Labiatae Lacandoniaceae Lachenaliaceae Lacistemataceae Lactoridaceae Lactucaceae

Lagerstroemiaceaee LamiaceaeLampsanaceae Lanariaceae Langsdorffiaceae Lantan

aceae Lapageriaceae Lardizabalaceae Lasiopetalaceae LauraceaeLawsoniaceae Laxm

anniaceae Lecythidaceae Ledaceae Ledocarpaceae Leeaceae Leguminosae Leitneriac

eae LemnaceaeLennoaceae Lentibulariaceae Lentiscaceae Leoniaceae Leonticaceae

Lepidobotryaceae LepidocarpaceaeLepidocaryaceae Lepidocerataceae Leptaulaceae

Leptospermaceae Lepturaceae Lepuropetalaceae LeuctenbergiaceaeLibocedraceae

Licaniaceae Ligustraceae Lilaeaceae Liliaceae Limeaceae Limnanthaceae Limnocharit

aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace

ae Linderniaceae Lindleyaceae Linnaeaceae LippayaceaeLiriaceae Liriodendraceae Li

ssocarpaceae Littorellaceae Loasaceae Lobeliaceaee Loganiaceae LomandraceaeLoni

ceraceae Lopeziaceae Lophiocarpaceae Lophiolaceae Lophiraceae Lophophytaceae L

ophopyxidaceae LoranthaceaeLotaceae Lowiaceae Lupulaceaee Luxemburgiaceae Lu

zuriagaceae Lychnidaceae Lyciaceae LyginiaceaeLygodisodeaceae Lysimachiaceae Ly

thraceae

Macarisiaceae Mackinlayaceae Madiaceae Maesaceae Magnoliaceae Malaceae Male

sherbiaceae MalortieaceaeMalpighiaceae Malvaceae Mangiaceae Manicariaceae Ma

paniaceae Marantaceae Marathraceae MarcgraviaceaeMartyniaceae Mastixiaceae

Matricariaceae Maundiaceae Mayacaceae Medeolaceae Medusagynaceae Peridiscac

eaeMelaleucaceae Melampyraceae Melanophyllaceae Melanthiaceae Melastomatac

eae Meliaceae Melianthaceae MelicaceaeMeliosmaceae Melittidaceae Melochiacea

e Memecylaceae Mendonciaceae Menispermaceae Menthaceae MenyanthaceaeMe

nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

Nypaceae Nyssaceae

Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

ceae Papilionaceae Pappophoraceae PapyraceaeParacryphiaceae Parasitaxaceae Par

ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

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hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

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cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

50

ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 15: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

15

Platanaceae - the choices made here follow common usage eg as in textbooks like

Judd et al (2007) and Simpson (2006) and particularly in the new edition of

Mabberleys The Plant Book (Mabberley 2008) This latter is itself an attempt to reflect a

consensus the result of taking the opinions of botanists at several meetings A largely

similar consensus is reflected in the APG III classification (APG 2009) where you will

see that alternative classifications have been dispensed with For many people the

existence of such alternative classifications will simply confuse so agreement over

which groupings to use when alternatives are permitted and then dispensing with the

alternatives that are not commonly used seems reasonable I am making the few

changes that are needed to bring the classification here in line with these others but

only gradually since there is not enough time in each day

As already mentioned higher-level relationships in general and the composition of

orders in particular have always presented something of a challenge to systematists

(eg Davis amp Heywood 1963) That being said the composition of some clades like

Apiales Crossosomatales and Pandanales is decidedly unexpected However it is

interesting to see that these higher level clades are generally accepted even in works

with different classificatory philosophies (eg Heywood et al 2007) For clades like

Malpighiales many of the family groupings within Asparagales etc attempts to find

distinctive characters have largely failed (but see Endress amp Matthews 2006a also

the Apomorphies page here) Interestingly as with families some groupings suggested

by molecular studies are supported by morphological andor chemical characters that

have long been known sometimes for over a hundred years the relationship between

Pittosporaceae and ApiaceaeAraliaceae are a case in point (Hegnauer 1969b and

references) As our knowledge of morphology and chemistry improves we can hope for

improvements in the characterisations of clades at all levels

In many cases the new family limits of the Angiosperm Phylogeny Group (see

APG 1999 2003) are not really controversial although changes from the limits

commonly accepted only a decade ago are sometimes quite dramatic (eg Wagenitz

1997) Thus the split of the old Saxifragaceae sl is necessitated by its extreme

polyphyly as also with Icacinaceaesl and Cornaceae sl However the limits and

relationships of such groups had long been considered to be unsatisfactory but until

recently there had been no real reason to prefer one proposed arrangement over

another It is generally accepted that the limits of Lamiaceae and Verbenaceae have to

be redrawn although there is as yet no compelling evidence that the redrawn taxa are

not sister taxa (there is no evidence that they are) But whatever their relationships the

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

16

content of the clades has changed considerably and incidentally they are now easier to

identify than before the decision to recognise two families is not difficult The same is

true for Salicaceae and Achariaceae (Malpighiales) two previously small families that

have received the bulk of the old Flacourtiaceae Clade and hence taxon limits remain

difficult around Theaceae (Ericalessee alsoSladeniaceae Pentaphylacaceae)

and Euphorbiaceae (Malpighiales see

also Phyllanthaceae Picrodendraceae Putranjivaceae etc) Here current groupings

may not represent quite such dramatic changes in our understanding of relationships

Indeed parts of the old Euphorbiaceae that were separated may yet go back together

particularly Phyllanthaceae and Picrodendraceae (see Wurdack et al 2004 Davis et al

2005) even if Theaceae should indeed be dismembered (Geuten et al 2004) However

given that there is no molecular evidence that currently warrants combining all the

segregates of Euphorbiaceae and even if some do come together the clades the

families represent suggest novel groupings not recognised in current classifications

Furthermore Rafflesiaceae appear to be embedded within Euphorbiaceae s str (Davis

et al 2007) so the family is divided Of course maintaining even a moderately narrowly-

circumscribed Euphorbiaceae would entail reducing the iconic Rafflesiaceae to

synonymy Relationships in core Caryophyllales especially

around Phytolaccaceae andNyctaginaceae and also Portulacaceae are incompletely

understood and refashioning of taxon limits will doubtless be needed as cladistic

relationships become apparent (eg see Nyffeler 2007 Ogburn 2007a b) Some

groupings in the old Icacinaceaeand Olacaceae also remain unclear

The discovery of the relationships of parasitic and aquatic groups have presented a

particular challenge to systematists Morphologically some of these plants are so highly

modified that interpretation of the plant body in conventional terms is difficult or even

impossible Thus parasitic groups (broad-leaved angiosperms only) such as Rafflesiaceae

are hard to place since both the vegetative body and the flowers are changed almost

beyond recognition (flowers of rafflesiaceae are a good example of gigantism - Davis et

al 2007) Furthermore plastid gene sequences may be difficult or impossible to obtain

the chloroplast DNA in particular being highly degraded and the rate of molecular

change in general may be high so the problem of long-branch attraction is serious (eg

Duff amp Nickrent 1997 Nickrent et al 1998 Caddick et al 2002a G Petersen et al

2006b) Echlorophyllous sapromycoheterotrophic taxa mostly known from monocots

present similar problems However progress is being made placements for

Apodanthaceae Rafflesiaceae Mitrastemonaceae Cytinaceae and Cynomoriaceae have

recently been suggested (Barkman et al 2004 2007 Davis amp Wurdack 2004 Nickrent et

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

17

al 2004 Davis et al 2007) it seems likely that Burmanniaceae sl are polyphyletic but

both parts are in Dioscoreales (Merckx et al 2006) and relationships within the largely

hemiparasitic Santalales are also gradually being clarified (Maleacutecot 2002) The inclusion

of parasitic taxa in general molecular analyses can cause conniptions (eg Nickrent et al

2004 Davis et al 2004 Chase et al 2006 G Petersen et al 2006b) and there can be

horizontal transmission of genes (eg Davis amp Wurdack 2005 Vitaceae to Rafflesiaceae

Barkman et al 2007 the mitochondrial atp1 gene commonly moves)

Similarly in water plants neither vessels in particular nor much xylem in general is

needed leaves are highly modified and water-mediated pollination if adopted may

well be associated with major changes in floral morphology Here too recent molecular

studies suggest that aquatic groups with hitherto problematic relationships may find

homes Thus Podostemaceae are close to Clusiaceae and Calophyllaceae and are sister

to Hypericaceae (Malpighiales Kita amp Kato 2001) Hydatellaceae which used to be in

Poales are part of Nymphaeales (Saarela et al 2007) and Hydrostachyaceae may be

close to Hydrangeaceae (Cornales Xiang et al 2002) or perhaps should be placed in

Lamiales note that in the first two cases in particular there are morphological and

chemical features that support such a move If Podostemaceae are indeed close to

Clusiaceae I look forward to seeing hypotheses to explain how the dramatic changes in

the vegetative body that have made Podostemaceae so problematic for generations of

systematists took place That conventional wisdom has trouble in understanding or

explaining how the morphologies of groups like Clusiaceae and Podostemaceae can be

related is largely a problem with conventional wisdom Ceratophyllaceae seem to be

finding a position as sister to eudicots (see Moore et al 2007) but their morphology is

so derived that there is no morphological evidence of which I am aware for this

relationship There are also similar although less extreme morphological problems with

many plants pollinated by wind

In such situations especially with some water plants mycoheterotrophs and

parasites a variety of characters may be affected and particularly under such

circumstances groups have tended to be formed using a few or even only a single

character that seems to provide evidence of relationships Add to this the tendency to

weight some characters particularly strongly on a priori grounds and the result was

either the recognition of conglomerate taxa such as Amentiferae which now appear to

be highly polyphyletic or the segregation of families like Plantaginaceae s str (now

much expanded) and Leitneriaceae (now in Simaroubaceae) - all of these are more or

less wind pollinated

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

18

Even although relationships of many parasitic and aquatic groups are being

clarified and at least some of the proposed phylogenetic positions seem to be well

supported it is indeed often still very difficult to understand how such groups have

evolved from their more morphologically conventional relatives However we have had

all sorts of preconceptions as to how characters should evolve Thus we used to

assume that features like highly scalariform vessel perforation plates or the complete

absence of vessels or a flower with an androecium that had many stamens a superior

ovary or separate petals were necessarily plesiomorphic or primitive and conversely

a simple perforation plate and the presence of vessels an androecium with few

stamens inferior ovary or petals that were connnate were almost necessarily

apomorphic or advanced (but cf eg Stebbins 1951) Such assumptions are incorrect

(eg Soltis et al 2005b) Carpels may become secondarily free carpels may fail to close

the seeds then developing outside the confines of the carpel as in some Aspagaraceae-

Nolinoideae Violaceae Berberidaceae Malvaceae-Sterculioideae etc) in Peliosanthes

teta perhaps the only species in Peliosanthes(Asparagaceae-Nolinoideae) the ovary

varies from superior to inferior (Jessop 1976 species limits here need close

investigation see also Kuzoff et al 2001 and Soltis amp Hufford 2002 Saxifragaceae

Apiales etc) many-seeded carpels can evolve from few-seeded carpels

(Razafimandimbison et al 2008) monoecy may be derived from dioecy (Schaefer amp

Renner 2010 and references) Classic studies such as those by Babcock (eg 1947)

on Crepis that assumed that evolution - in this case of the karyotype in particular - was

unidirectional have needed comprehensive re-evaluation (Enke amp Gemeinholzer 2008)

Most if not all characters have reversed andor evolved in parallel as is clear in the

discussion of ovary position here especially in Poales Saxifragales and Asparagales and

also in comments on such morphologically distinctive taxa (when compared with their

immediate relatives) such as Menyanthaceae and especially Pittosporaceae Parallel

evolution may occur even at the level of amino acid substitution as in the independent

acquisition of the phosphoenolpyruvate carboxylase (pepC) gene in C4 photosynthesis in

grasses (Christin et al 2007b see also Blaumlsing et al 2000)

Thinking of how such characters have been used in the past in classifications at the

generic level the results have been similar to those just discussed above for parasitic

plants and others Relying too much on animal pollination syndromes to mark generic

boundaries has all too often led to taxa that are highly unsatisfactory phylogenetically

(see eg Acanthaceae Bignoniaceae Campanulaceae Ericaceae Lamiaceae

Orchidaceae) and over-reliance on characters of fruit and seed (see eg Brassicaceae

and Apiaceae) has also led to unsatisfactory generic limits Again the more general

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

19

problem is the use of one or a very few characters that have been weighted a priori to

structure classifications (see also Garciacutea et al 2009) although it is perhaps not

surprising that such an approach sometimes results in the delineation of taxa that have

indeed turned out to be quite distinct (cf eg Burtt Astragalus versus Oxytropis)

Taxa recognized here are monophyletic yet many are polythetic at the

morphological level that is they lack unique features characterizing (defining) all and

only members of that family they can be recognised phenetically only by the unique

combinations of characters that they posess This is the result of evolution any

synapomorphy characterizing a taxon may be lost or modified beyond easy recognition

in some of its members or the synapomorphy may appear to be identical to a feature

that has evolved in parallel in a quite unrelated plant That plant groups are polythetic is

almost as much a feature of monophyletic taxa as the paraphyletic taxa common in

evolutionary classifications Some families now include substantial variation as

phenetically distinct derived groups are placed in their proper phylogenetic position -

examples are Ericaceae which include the erstwhile Empetraceae a wind-pollinated

group that is florally very different and the overwhelmingly large-flowered and animal-

pollinated Plantaginaceae which now include several derived small-flowered aquatic

and wind-pollinated groups that were previously placed in separate families

Thus some families as delimited here may not be easy to recognize However

remember that detecting relationships - use whatever characters you can even if they

are not obvious - and naming a plant - focus on easy-to-see characters that may not

reflect relationships - are quite different problems Taxa although natural may not be

readily recognizable indeed it was in exactly this context that Lamarck worked out the

basic principles of writing dichotomous keys in 1778 (see Scharf 2007 for keys and the

like) Of course Lamarcks idea of nature was very different from ours - he thought that

there was some kind of continuum of form on which living organisms were to be

situated with no real gaps anywhere - but this meant that his genera (for example)

might well not be sharply distinct from each other even if each were part of the real

continuum that was life Lamarcks separation of the establishment of relationships and

groupings in a system and identification of the plants in that system have been widely

accepted (but cf eg Godfray amp Knapp 2004 many of the contributors to Heywood et

al 2007) Perhaps the best way of identifying plants at the family level is by well-made

multiple access keys as in Watson and Dallwitz (1992a onwards family limits there may

differ substantially from those adopted here) Multiple access keys free users from the

constraints of dichotomous keys in which particular characters are needed at each step

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

20

of the identification process before there can be further progress Instead those

characters that happen to be evident on a specimen can be used in whatever order is

convenient when linked to illustrations glossaries etc their power is enormous (see

Dallwitz et al 2000 [2006] for the principles underlying their construction and use)

Nevertheless dichotomous keys such as those of Hutchinson (1973) and Franz Thonner

(Geesink et al 1981) have their uses Of course taxa coming out adjacent in keys may

well not be at all related

(When identifying large numbers of plants even more efficient than either style of

identification and certainly lots more fun is sight identification Unless you have a

photographic memory you have to build up your knowledge of comparative plant

morphology - on which the ability to make accurate identifications depends - by

repeated observation When faced with an unknown plant I always look for leaf teeth

and stipules distinctive hair types especially stellate and T-shaped hairs glands and

punctations of any sort the presence of latex or other exudate and check leaf insertion

smelling crushed leaves can also be helpful In this context nodal anatomy can usually

be checked using a razor and a hand lens or even simply carefully examining leaf scars

The short paragraphs added after most families may help in confirming familial

identifications)

For the record and for the little that it is worth there are 4 orders and 13 families

of gymnosperms characterised on these pages and together they include some 82

genera and 947 species For angiosperms comparable figures are 56 orders 445

families 13208 genera and 261750 species (of which monocots include 11 orders 89

families 2759 genera and 52760 species) Note however that higher mathematics

was never my strong point and anyway these are pretty meaningless figures even for

species which many (but not all) might concede smacked slightly more of reality than

other taxa estimates range as high as 422000 (Govaerts 2001) Furthermore numbers

of genera and species change daily Nevertheless as emphasized here families are

useful in teaching we as a community can ensure that their limits remain largely stable

and by concentrating on relatively few of them one can gain some familiarity with much

of the worlds flora For summaries of the sizes of orders and families and of the general

arrangement followed in these pages see also the Statistics page itself

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

21

A PHYLOGENETIC TREE OF SEED PLANTS (THE MAIN TREE)

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

22

The Main Tree above is a conservative summary of well supported relationships

between the major clades of extant seed plants Reading - interpreting and

understanding - such a phylogenetic tree is not necessarily easy so reading a tree may

require a bit of practice OHara (eg 1988 1992) provides an accessible introduction to

thinking and talking about history in the context of looking at phylogenetic trees More

recently Krell and Cranston (2004) Crisp and Cook (2005) and others have emphasized

how careful one must be when interpreting and talking about evolution in ladderized

trees in particular and phylogenetic trees in general Gregory (2008) provides a

particularly detailed discussion of this problem In ladderized trees the smaller (in terms

of numbers of terminals) sister taxon is consistently shown on the same side at every

node thus the trees tend to be pectinate like the teeth of a comb however the

horizontal axis that is as a result so evident has no polarity or particular direction The

use of the adjective basal when discussing phylogenetic trees is especially dangerous

(see also Wojciechowski et al 2004 D Soltis et al 2005b) When I use the term and the

context is not otherwise clear I am referring to the pectinations at the base of a

ladderised tree Remember that when talking about sister taxa one can never be basal

to the other or older or younger although members of one can be more derived - in the

sense of having more apomorphies - than the other (but even then one has to be

careful) The words primitive and advanced and lower and higher should

practically never be used especially when talking about taxa Amborellaceae and

Pinaceae are sister to all other flowering plants and Pinales and are basal to the crown

groups of all other flowering plantsPinales respectively but that does not mean they

are thereby primitive Pinaceae in particular have numerous apomorphies The word

plesiomorphic is far less loaded than primitive and can be used to talk about

individual characters

Most trees here have been more or less ladderized as is the one above The

ladderization may be imperfect for example you can see that on the Main Tree

asterids with ten orders follow rosids and relatives which have seventeen although

this is in part because of the currently unresolved nature of relationships between the

main clades of core eudicots (this is likely to change soon) The Main Tree could have

been drawn with Amborella Acorus or a host of other taxa at the far right without

offending any relationships phylogenetic trees are like mobiles the only fixed points

being the nodes Despite the problems just mentioned pectination interpreted

carefully has its value and it is not an arbitrary process As one reads the terminals of a

pectinate tree from left to right adjacent terminals will be separated by apomorphies

that will cumulate in an appropriate fashion When thinking of a book or a herbarium

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

23

sequence (see Haston et al 2007) this is of some value Nymphaeales and

Austrobaileyales are here adjacent on the tree but they could be separated by

hundreds of families in the sequence merely by rotating only the node from which

Nymphaeales and their sister group (all other angiosperms minus about 75 species)

arise If adjacent in a book or herbarium or as in these summary pages then it is

relatively easy to relate their characters both apomorphic and plesiomorphic but if

separated by hundreds of pages or two floors in a large building then it is less easy to

get anything from the sequence (Since all orders in the pages of the main part of this

site are preceded by the apomorphies of all nodes immediately below them in the seed

plant phylogeny and because of the linkages that have been built in to the site this

problem is much less serious there there is no sequence) So the particular sequence of

all seed plants that is based on a phylogenetic classification but is to be used in herbaria

or any other place where taxa have to be arranged linearly as in this part of APweb

will be one that maximizes the number of taxa that are both successive branches of the

tree and placed successively in the sequence Since specimens are generally filed under

families the outline of a new family sequence for arranging herbaria and books can now

be suggested (Haston et al 2007) and such a standardized sequence will further help

teaching and learning about plants

ON THINKING ABOUT APOMORPHIES

Identifying apomorphies is important because understanding the evolution of

morphology in the broad sense - ie including anatomy chemistry etc - in the context

of diversification is one of our major goals For identifying apomorphies several

preconditions must be met One needs to have an accurate well supported phylogeny

one has to have examined the right taxa both from the point of view of morphology and

molecules one has to have coded the characters right (ie delimited states

appropriately) and one has to use an appropriate model of evolution when fitting the

variation to the tree (see eg Omland 1999 Stevens 2006b) If the distributions of

apomorphies below are compared with those in other studies (eg Turgeon et al 2001

Bremer et al 2001 Endress 2001 Albach et al 2001a Judd amp Olmstead 2004 D Soltis

et al 2005b Zhang et al 2006 Judd et al 2007) the reader may find a number of

differences Although I have integrated such studies as far as is possible there are five

reasons that link to the preconditions just mentioned why there may be differences

where features are placed on trees

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

24

1 I may not have found all the information about a particular character there may be

disagreement over its interpretation or I have added information

2 The sampling of nearly all molecular studies is very incomplete (see Salisbury amp Kim

2001 for problems caused by sampling) indeed it is commonplace to decry the

incompleteness of molecular sampling But not only is the sampling in molecular studies

often less than we might wish that of the morphological and chemical characters whose

evolution we are interested in understanding is also often very poor So for many

anatomical chemical and embryological characters that are confidently said to

characterise families and other groups we all too often have no idea if those characters

are applicable to the whole clade or just to a subgroup within that clade However as

our sampling improves we can locate changes on the tree more precisely if in

unexpected positions Thus is is sometimes suggested that Ericaceae have ellagic acid

(Soltis et al 2005b) in fact the little evidence we have indicates that only a small clade

of some 80 species has ellagic acid and the rest of the family - itself alone about one

third of all Ericales - does not Similarly Ericaceae are noted for their

ectendomycorrhizal associations but Enkianthus sister to all other Ericaceae

apparently lacks these (Abe 2005) Thus these mycorrhizae and their associated

distinctive hair roots are unlikely to be apomorphies of Ericaceae - however it appears

that other families near Ericaceae may also have ectendomycorrhizae (see Asai 1934)

3 I am fitting characters to a very conservative tree with many polytomies although the

nodes that are utilised are for the most part strongly supported polytomies make the

optimisation of characters that is the assigment of character state change to a

particular node on the tree notably difficult (eg Madison amp Madison 2002) In nearly

all studies of the evolution of characters distributions of characters are optimised on a

more or less fully resolved tree and the construction of supertrees may yield yet more

detailed hypotheses of relationships (for literature on supertrees see Cotton amp

Wilkinson 2007 2008) Of course some nodes on such fully resolved trees andor

supertrees may have little support and optimisations of characters on such trees may

carry correspondingly little conviction Even parts of some of the trees used here have

poor support eg relationships within aquatic Alismatales etc although I indicate

when this is the case

4 Exactly how one goes about optimising a character on a tree is critically important

Even using simple parsimony optimisations (ACCTRAN or DELTRAN ACCelerated

TRANsitions or DELayed TRANsitions) the position of synapomorphies on trees - and

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

25

hence our ideas of evolution - may differ and this is still more true if one uses maximum

likelihood or Bayesian methods and makes apparently reasonable suggestions about

weighting gains over losses (eg Donoghue amp Ackerley 1996 Cunningham et al 1998

Omland 1997 1999 Ree amp Donoghue 1999 Polly 2001 Webster amp Purvis 2001

Ronquist 2004 Crisp amp Cook 2005) Sannier et al (2007) show how in in Arecaceae that

where on a phylogenetic tree one might peg changes in microsporogenesis will depend

on the methods one uses to do this and Pedersen et al (2007) discuss the sometimes

very substantial effect of node support on the posterior probabilities of ancestral

character states Here I use parsimony optimization not always as explicit as it might be

but I have often indicated where there are particularly important uncertainties as to the

positions of particular character changes on the tree

5 Finally although I have paid quite a lot of attention to the delimitation of the

character states that make up all the characterizations I have not spent enough time on

this critical operation If we are interested in understanding evolution then fitting the

basic variation - not character states - to a tree in principle allows greater flexibility in

understanding morphology in the context of local phylogenies (see also Stevens 2000

Endress 2005c) However many character states used here are delimited globally that

is they are circumscribed in the context of the variation shown by individual characters

across all angiosperms andor in the context of classic ideas of character evolution

Character states often have arbitrary limits and serve best to communicate

information whether they are in fact suitable for either phylogenetic analysis or

understanding evolution are separate issues Studies have rather unsurprisingly

perhaps but importantly shown that dividing the one character into different sets of

states may yield differing ideas of evolution of that character (eg Lamb Frye amp Kron

2003 Hibbett 2004) When looking at trees on which character states are optimised

one should bear in mind the problems surrounding the delimitation of states (eg

Stevens 2000 2006b) and the danger of using pollen or other types - constructs

based on many characters that vary independently but which effectively get lost in

these types thus Blackmore et al (2009) decomposed the pollen types in Asteraceae

into 52 characters

I use Remanes three main criteria of homology or better similarity when

determining the basic similarity of structures on different organisms (see Remane 1952)

These criteria are special properties position and intermediates Special properties

include anatomical or chemical characters gene expression data etc Position refers

to the position of an organ with respect to landmarks on the plant Although plants are

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

26

plants and landmarks are not as fixed as one might like trying to work out the relative

positions of parts is a good way of understanding morphology so long as one does not

try to out-do Procrustes Intermediates include intermediates found on the same

plant or on different plants Intermediates may observed during development or be

adult structures Thus one may find changes during development which suggest what

the nature of a particular structure is and very different stuctures on different plants

may turn out to be similar early in development Or one can simply compare different

taxa and see that structures that appear to be very different when comparing two

different taxa can be linked morphologically by looking at other taxa

Nevertheless the use of these criteria may not yield an unambiguous answer as to

what a structure is even given a solid phylogeny and an improved understanding of

development (see Jaramillo amp Kramer 2007 for a useful discussion) As Endress (2005c)

observed a number of features - position function development shape anatomy

histology gene activity and relationships to other taxa that clearly have petals - can be

used to distinguish a petal (for example) from other floral structures if a petal does not

have one of these features is it thereby not a petal Thus Maturen et al (2005) recently

found that floral organ diversity genes (B and C) were expressed in the large white

inflorescence bracts of Cornus (see also Costa et al 2005) Peney et al (2005) noted that

not all monosulcate pollen grains in monocots have the same developmental pathway

and that as a result such pollen might not have the same ancestral state Reeves and

Olmstead (2003) suggested that the genetic mechanisms causing monosymmetry in

Lamiales and Solanales were different and Serna and Martin (2006) described similar

problems with the development of hairs in Arabidopsis when compared with that of

hairs in Antirrhinum and Solanaceae Indeed as one perhaps might expect delimitation

of states and characters does not necessarily become easier with increasing knowledge

of development etc Thus Buzgo et al (2004) Matthews and Endress (2005) and others

have shown how hard it can be to distinguish between eg prophylls and other floral

structures as their behaviour is studied during the course of the development of the

flower Similarly at what concentration is a particular secondary metabolite deemed to

be present (Waterman (2007)

The validity of the approach used here that of fitting morphological variation to a

largely molecular-based tree may be questioned However I think it rather unlikely that

well-supported molecular branches will be overturned by morphological data Indeed

analyses of morphological data alone do provide support for many of the clades evident

in molecule-only analyses and analyses with morphological and molecular data

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

27

together may lead to increased support for clades (eg Hufford 1992 Nandi et al 1998

[but adding morphological data reduces support for a number of critical clades too]

Doyle amp Endress 2000 however in none of these papers is the use of morphology

without ambiguity) It is unfortunately clear that the use of morphology alone may not

suggest problems in the phylogenetic placement of taxa that later turn out to have been

wrongly included (eg Zhang et al 1992) Thus I have been wary of putting much weight

on clades that have only morphological support but note that I have treated molecule-

based clades with low bootstrap or jacknife support values (esp below 70) or low

posterior probabilies (below 095) likewise Although I may have been mistaken in

placing so much emphasis on molecular data in terms of providing the basic

phylogenetic framework for angiosperms morphological and molecular data are only

very rarely in irreconcilably strong conflict There are indeed a few places where the

conflict seems extreme These include the relative positions of the Monimiaceae and

Hernandiaceae (Laurales) the position of Hanguanaceae (Commelinales [as here] or

Zingiberales) and of Triplostegia (is it in Dipsacaceae or Valerianaceae - see

Dipsacales) Fossils are unlikely to affect the topologies of the trees presented here but

see below for their importance in understanding morphological evolution in general and

the evolution of angiosperms in particular Nevertheless some largely reject the idea

that trees based on molecular data alone can recover phylogenetic relationships

especially when branching points are old and prefer to used trees based on analysis of

morphological data including those taken from fossils (Hilton amp Bateman 2006 Farjon

2007)

All in all however the extent of the congruence between morphological and

molecular data is impressive and heartening and many clades can be characterised

morphologically It seemed in 1998 that there were no unambiguous morphological

synapomorphies for angiosperm orders (K Bremer 2000) and this is still true if by

unambiguous is meant non-homoplasious However many orders and other clades

have synapomorphies even if these may be indistinguishable at least at the current

level of morphological and developmental knowledge from parallel occurences

elsewhere As our morphological knowledge increases so too does the number of

apomorphies

Endress and Matthews (2006a) emphasize the importance of tendencies and

developmental constraints when thinking about characters of clades within the rosids -

many characters or character combinations occur in a rather sporadic fashion within

the clade and apparently notably less frequently outside it There are a number of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

28

examples of tendencies here and in the individual order pages such as the features

enclosed by parentheses in groups above the level of family eg see (cuticular waxes

as aggregated rodlets) for the commelinids and the discussion of the distribution of

polyandry that is flowers with numerous stamens in the asterid I + II groups etc

Tendencies may involve more than single characters When thinking of the

associations of features like integument thickness and vascularization of the integument

(Endress amp Matthews 2006a) one can imagine fairly simple developmental

preconditions being operative As Endress and Matthews (2006a) note it is difficult to

imagine vascular tissue developing in an integument only two cells thick Changes here

would be loosely correlated if morphologically linked However the strongly correlated

changes noted by Givnish et al (2005) are ecologically linked but are presumably

morphologicallydevelopmentally independent When there seem to be characters

evolving more or less together tests can be carried out to see if the changes are

concentrated on certain branches of the tree (eg Maddison 1990 Sanderson 1991

Maddison amp Maddison 2000) Such tests have rarely been carried out at higher levels in

angiosperms

One of the most striking examples of a tendency is the distribution of N-fixation

restricted as it is to a monophyletic group of four clades although it has arisen seven (or

perhaps several more) times independently within the N-fixing clade and several

members of two quite different kinds of bacteria are involved (eg Clawson et al 2004

Elliott et al 2007 Sprent amp James 2007) Other examples of work that bears on the issue

of tendencies include the findings that flowers of polysymmetrical Arabidopsis have

genes like TCP1 that are expressed asymmetrically during early development and TCP1

is a probable orthologue of the well-known CYC gene of Antirrhinum that is involved in

the development of monosymmetric slowers there (Cubas et al 2001 Costa et al 2005

etc) Parallelism might build on this underlying morphologically cryptic monosymmetry

even if details of the genetic mechanisms causing the monosymmetry evident in

particular groups may be different (Reeves amp Olmstead 2003 Cubas 2004 see above)

Similarly there have been several recent suggestions that the capability to synthesise a

particular metabolite may be switched off but not lost and so can sometimes be

reacquired (eg Wink amp Witte 1983 Wink 2003 Liscombe et al 2005 Larsson 2007

Waterman 2007) Hence perhaps the rather spotty distribution of many secondary

metabolites like ellagic acid the indole alkaloid camptothecin iridoids etc when

considered in the context of phylogenies As a non-botanical example - but a rather nice

one - Salwini-Plawen and Mayr (1961) suggested some time ago that there has been

considerable parallelism (40-65 or more independent origins) in the evolution of eyes in

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

29

metazoans However Pax 6 seems to be a master control gene that is involved in eye

formation perhaps throughout all bilateralians (eg Gehring amp Ikeo 1999 Erwin amp

Davidson 2002) which changes the nature of the problem Note however that in

general our invocation of evolutionary tendencies to explain the patterns of

distributions of characters is really an admission that we do not understand those

patterns

As we find out more about variation we will probably find fewer and fewer features

constant throughout a group Most unqualified statements of presence and absence

should properly be qualified as usually present or usually absent if one is thinking of

the characterisations on the order pages as encompassing the total variation within a

clade Thus Pistia alone among monocots as so far known has sieve tube plastids with

starch grains not protein crystals However this does not affect the fact that sieve tube

plastids with cuneate protein crystals and lacking starch grains are an apomorphy for

monocots Furthermore fossils in a number of cases suggest character combinations

unknown in extant taxa as may be seen in the discussions of Fagaceae Platanaceae

Iteaceae Calycanthaceae etc Confusing the issue there may be questions as to where

exactly on the tree a particular fossil is to be placed (eg see Nymphaeaceae

Calycanthaceae Archaefructus etc)

To summarize given our current understandings of both phylogenies and

characters evolution of some characters in which we are interested seems very labile

(see eg D Soltis et al 2005b Endress and Matthews 2006a Stevens 2006b Ekman et

al 2008 for a good recent study) and I have been cautious when talking about character

evolution Much effort must continue to be spent in summarizing characters of well-

established clades at all levels providing features by which they may be recognized and

signaling synapomorphies Remember that (1) the basic morphological anatomical and

chemical knowledge of many critical taxa is woefully incomplete (2) different

assumptions about character evolution may greatly affect the position of

synapomorphies on trees (3) in many cases relationships within and between many

groups are too uncertain at present to worry very much about synapomorphies and

(4) we must be clear about what we do and do not not know As mentioned above I

have indicated in a number of places where there are particularly important

uncertainties as to where characters should be placed on the tree Nevertheless it is a

relatively easy matter to update notes such as these and it can be a simple matter to

incorporate new data on characters that have never before been considered in the

context of a tree Much basic - and unfortunately perhaps unfashionable - work must be

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

30

carried to clarify the distribution of morphological anatomical and chemical characters

for examples of what can to be done see the work of P K Endress and collaborators

(floral morphology and development) and S R Jensen and collaborators (iridoids) But

acquiring information about nodal anatomy cell and tissue distribution nodal anatomy

and the like is not difficult at all and should be generally encouraged

But all this is merely a necesary prelude to the understanding of evolution For this

one needs to know a lot more including dating the phylogeny understanding the

function(s) of characters factoring in relevant aspects of the palaeoenvironment in

which an apomorphy first appeared etc

SUMMARY OF THE SYSTEM

Below is a formalised summary of the relationships within orders of the families of seed

plants There are a few families that are not recognised even as options in APG II and

vice versa as well as a few extra orders The families to be recognised in the most

recent edition of Mabberleys The Plant Book (Mabberley 2008) are also largely

consistent with those below But all differences are trivial and will - I hope - eventually

disappear

Square brackets - [] - enclose clades the plus sign - + - designates sister taxa a

comma - - denotes part of a polytomy and quotation marks - - denotes a

paraphyletic group

SEED PLANTS

GYMNOSPERMS

Cycadales

Cycadaceae + Zamiaceae

Ginkgoales

Ginkgoaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

31

Pinales

Pinaceae [[Araucariaceae [Podocarpaceae + Phyllocladaceae]] [Sciadopityaceae

[Taxaceae + Cupressaceae]]]

Gnetales

Ephedraceae [Gnetaceae + Welwitschiaceae]

ANGIOSPERMSFLOWERING PLANTS

Amborellales

Amborellaceae

Nymphaeales

Hydatellaceae [Cabombaceae + Nymphaeaceae]

Austrobaileyales

Austrobaileyaceae [Schisandraceae + Trimeniaceae]

Chloranthales

Chloranthaceae

MAGNOLIIDS

Magnoliales

Myristicaceae [Magnoliaceae [[Himantandraceae + Degeneriaceae] [Eupomatiaceae +

Annonaceae]]]

Laurales

Calycanthaceae [[Siparunaceae [Gomortegaceae + Atherospermataceae]] [Monimiaceae

[Hernandiaceae + Lauraceae]]]

Canellales

Canellaceae + Winteraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

32

Piperales

[Hydnoraceae + Aristolochiaceae] [Piperaceae + Saururaceae]

MONOCOTS

Acorales

Acoraceae

Alismatales

Araceae [Tofieldiaceae [[Alismataceae [Hydrocharitaceae + Butomaceae]]

[Scheuchzeriaceae [Aponogetonaceae [Juncaginaceae [Maundiaceae [[Posidoniaceae

[Ruppiaceae + Cymodoceaceae]] [Zosteraceae + Potamogetonaceae]]]]]]]

Petrosaviales

Petrosaviaceae

Dioscoreales

Nartheciaceae [[Taccaceae + Thismiaceae] [Burmanniaceae + Dioscoreaceae]]

Pandanales

Velloziaceae Triuridaceae Stemonaceae [Pandanaceae + Cyclanthaceae]

Liliales

Corsiaceae [Campynemataceae [Petermanniaceae [Colchicaceae + Alstroemeriaceae]]

Melanthiaceae [[Philesiaceae + Rhipogonaceae] [Smilacaceae + Liliaceae]]]

Asparagales

Orchidaceae [[Boryaceae [Blandfordiaceae [Lanariaceae [Asteliaceae + Hypoxidaceae]]]]

[[Ixioliriaceae + Tecophilaeaceae] [Doryanthaceae [Iridaceae [Xeronemataceae

[Xanthorrhoeaceae [Alliaceae + Asparagaceae]]]]]]]

COMMELINIDS

Unplaced

Dasypogonaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

33

Arecales

Arecaceae

Poales

[Typhaceae + Bromeliaceae] [Rapateaceae [[Thurniaceae [Juncaceae + Cyperaceae]]

[[Mayacaceae [Eriocaulaceae + Xyridaceae]] [Flagellariaceae [Anarthriaceae

[Centrolepidaceae + Restionaceae]]] [Joinvilleaceae [Ecdeiocoleaceae + Poaceae]]]]]]]

Commelinales

[Commelinaceae + Hanguanaceae] [Philydraceae [Haemodoraceae + Pontederiaceae]]

Zingiberales

Musaceae [Strelitziaceae + Lowiaceae] Heliconiaceae [[Cannaceae + Marantaceae]

[Costaceae + Zingiberaceae]]

Ceratophyllales

Ceratophyllaceae

EUDICOTS

Ranunculales

Eupteleaceae [Papaveraceae [[[Lardizabalaceae + Circaeasteraceae] [Menispermaceae

[Berberidaceae + Ranunculaceae]]]]

Sabiales

Sabiaceae

Proteales

Nelumbonaceae [Platanaceae + Proteaceae]

Trochodendrales

Trochodendraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

34

Buxales

Haptanthaceae [Buxaceae + Didymelaceae]

CORE EUDICOTS

Gunnerales

Gunneraceae + Myrothamnaceae

Dilleniales

Dilleniaceae

Saxifragales

Peridiscaceae [[Paeoniaceae [Altingiaceae [Hamamelidaceae [Cercidiphyllaceae +

Daphniphyllaceae]]]] [[Crassulaceae [Aphanopetalaceae [Tetracarpaeaceae

[Penthoraceae + Haloragaceae]]]] [Iteaceae [Grossulariaceae + Saxifragaceae]]]]

Cynomoriaceae unplaced

Vitales

Vitaceae

ROSIDS

FABIDROSID I

Zygophyllales

Krameriaceae + Zygophyllaceae

Celastrales

Lepidobotryaceae + Celastraceae

Oxalidales

Huaceae [[Connaraceae + Oxalidaceae] [Cunoniaceae [Elaeocarpaceae [Brunelliaceae +

Cephalotaceae]]]]

Malpighiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

35

[Achariaceae [Goupiaceae [Violaceae + Passifloraceae] [Lacistemataceae Salicaceae]]

[Lophopyxidaceae Putranjivaceae] [Ctenolophonaceae [Erythroxylaceae +

Rhizophoraceae]] Linaceae Ixonanthaceae Humiriaceae Irvingiaceae

Centroplacaceae Pandaceae [[Bonnetiaceae + Clusiaceae] [Calophyllaceae

[Hypericaceae + Podostemaceae]]] [Malpighiaceae + Elatinaceae] Ochnaceae

[Peraceae [Rafflesiaceae + Euphorbiaceae]] [Phyllanthaceae + Picrodendraceae]

[Balanopaceae [[Trigoniaceae + Dichapetalaceae] [Chrysobalanaceae +

Euphroniaceae]]] Caryocaraceae

N-FIXING CLADE

Fabales

Quillajaceae [Fabaceae [Polygalaceae + Surianaceae]]

Rosales

Rosaceae [[Barbeyaceae [Elaeagnaceae Dirachmaceae Rhamnaceae]] [Ulmaceae

[Cannabaceae [Moraceae + Urticaceae]]]]

Cucurbitales

Anisophylleaceae [[Corynocarpaceae + Coriariaceae] [Cucurbitaceae [Tetramelaceae

[Datiscaceae + Begoniaceae]]]] Apodanthaceae

Fagales

Nothofagaceae [Fagaceae [[Myricaceae + Juglandaceae] [Casuarinaceae

[Ticodendraceae + Betulaceae]]]]

MALVIDROSID II

pgtGeraniales

Geraniaceae [[Melianthaceae + Francoaceae] [Vivianaceae + Ledocarpaceae]]

Myrtales

Combretaceae [[Onagraceae + Lythraceae] [[Vochysiaceae + Myrtaceae]

[Melastomataceae [Crypteroniaceae [Alzateaceae + Penaeaceae]]]]]

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

36

Crossosomatales

[Staphyleaceae [Guamatelaceae [Crossosomataceae + Stachyuraceae]]] [Aphloiaceae

[Geissolomataceae + Strasburgeriaceae]]

Picramniales

Picramniaceae

Sapindales

Biebersteiniaceae Nitrariaceae [Kirkiaceae [Anacardiaceae + Burseraceae]]

[Sapindaceae [Simaroubaceae Rutaceae Meliaceae]]

Huerteales

Gerradinaceae [Dipentodontaceae + Tapisciaceae]

Malvales

Neuradaceae [Thymelaeaceae [Sphaerosepalaceae Bixaceae [Cistaceae

[Sarcolaenaceae + Dipterocarpaceae]] [Cytinaceae + Muntingiaceae] Malvaceae]]

Brassicales

[Akaniaceae + Tropaeolaceae] [[Moringaceae + Caricaceae] [Setchellanthaceae

[Limnanthaceae [[Koeberliniaceae [Bataceae + Salvadoraceae]] [Emblingiaceae

[Pentadiplandraceae [Gyrostemonaceae Resedaceae Stixis and relatives] Tovariaceae

[Capparaceae [Cleomaceae + Brassicaceae]]]]]]]]

Berberidopsidales

Aextoxicaceae + Berberidopsidaceae

Santalales

Erythropalaceae Olacaceae [[Loranthaceae [Misodendraceae + Schoepfiaceae]]

[Opiliaceae + Santalaceae]] Balanophoraceae unplaced

Caryophyllales

[[Droseraceae [Nepenthaceae [Drosophyllaceae [Ancistrocladaceae +

Dioncophylleaceae]]]]] [[Frankeniaceae + Tamaricaceae] [Polygonaceae +

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

37

Plumbaginaceae]]] [Rhabdodendraceae [Simmondsiaceae [[Asteropeiaceae +

Physenaceae] [[Caryophyllaceae [Achatocarpaceae + Amaranthaceae]]

[Stegnospermataceae [Limeaceae [[Lophocarpaceae [Barbeuiaceae [Aizoaceae

[Sarcobataceae Phytolaccaceae Nyctaginaceae]]]] [Molluginaceae [Halophytaceae

Montiaceae Basellaceae Didiereaceae [Talinaceae [Portulacaceae [Anacampseros etc +

Cactaceae]]]]]]]]]]]]]

ASTERIDS

Cornales

[Cornaceae + Nyssaceae] [Hydrangeaceae + Loasaceae] Hydrostachyaceae [Curtisiaceae

+ Grubbiaceae]

Ericales

[Balsaminaceae [Marcgraviaceae + Tetrameristaceae]] [[Polemoniaceae +

Fouquieraceae] Lecythidaceae [[Sladeniaceae + Pentaphylacaceae] [Sapotaceae

[Ebenaceae [Maesaceae [Theophrastaceae [Myrsinaceae + Primulaceae]]]]]

[Mitrastemonaceae Theaceae [Symplocaceae [Styracaceae + Diapensiaceae]]

[[Sarraceniaceae [Roridulaceae + Actinidiaceae] [Clethraceae [Cyrillaceae +

Ericaceae]]]]]]

LAMIIDASTERID I

Unplaced

Metteniusaceae Oncothecaceae Icacinaceae and some unplaced ex-Icacinaceae

Garryales

Garryaceae + Eucommiaceae

Unplaced

Boraginaceae Vahliaceae

Gentianales

Rubiaceae [Gentianaceae [Loganiaceae [Gelsemiaceae + Apocynaceae]]]

Lamiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

38

Plocospermataceae [Oleaceae [Tetrachondraceae [Calceolariaceae

[Gesneriaceae Peltanthera [Pedaliaceae Carlemanniaceae Martyniaceae Stilbaceae

Plantaginaceae Linderniaceae Bignoniaceae Verbenaceae Lamiaceae Paulowniaceae

Schlegeliaceae Thomandersiaceae Phrymaceae Rehmannia Acanthaceae

Scrophulariaceae Orobanchaceae Byblidaceae Lentibulariaceae]]]]]

Solanales

[Montiniaceae [Sphenocleaceae + Hydroleaceae]] [Convolvulaceae + Solanaceae]

CAMPANULIDASTERID II

Aquifoliales

[[Cardiopteridaceae + Stemonuraceae] [Aquifoliaceae [Helwingiaceae +

Phyllonomaceae]]

Asterales

[Rousseaceae + Campanulaceae] Pentaphragmataceae [[Alseuosmiaceae [Phellinaceae

+ Argophyllaceae]] Stylidiaceae [Menyanthaceae [Goodeniaceae [Calyceraceae +

Asteraceae]]]]

Escalloniales

Escalloniaceae

Bruniales

[Bruniaceae + Columelliaceae]

Apiales

Pennantiaceae [Torricelliaceae [Griseliniaceae [Pittosporaceae [Araliaceae

[Myodocarpaceae + Apiaceae]]]]]

Paracryphiales

Paracryphiaceae

Dipsacales

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Adoxaceae [Diervillaceae [Caprifoliaceae [Linnaeaceae [Morinaceae [Dipsacaceae +

Valerianaceae]]]]]

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MAIN GROUPINGS

Asterids commelinids core eudicots asterid 1 asterid

2 eudicots gymnosperms Magnoliophyta monocots N-fixing claderosids seed plants

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Abietales Acanthales Acerales Acorales Actinidiales Actinostrobales Adoxales Aescul

ales Agavales Aizoales AkanialesAlismatales Alliales Alseuosmiales Alstroemeriales

Altingiales Amaranthales Amaryllidales Amborellales AmbrosialesAmmiales Amomal

es Ancistrocladales Anisophylleales Annonales Anthobolales Apiales Apocynales Apo

nogetonalesAquifoliales Arales Araliales Aralidiales Araucariales Arecales Aristolochi

ales Athrotaxidales Asarales AsclepiadalesAsparagales Asphodelales Asparagales Ast

eliales Atriplicales Aucubales Austrobaileyales Avenales

Balanitales Balanopales Balanophorales Balsaminales Barbeyales Barclayales Batales

Begoniales BerberidalesBerberidopsidales Betulales Biebersteiniales Bignoniales Bix

ales Boraginales Brassicales Brexiales Bromeliales BrunialesBrunoniales Burmannial

es Burserales Butomales Buxales Byblidales

Cactales Callitrichales Calycanthales Calycerales Campanulales Campynematales Can

ellales Cannales CapparalesCaprifoliales Cardiopteridales Carduales Caricales Carle

manniales Caryophyllales Cassiales Casuarinales CelastralesCentrolepidales Cephalot

ales Cephalotaxales Ceratophyllales Cercidiphyllales Chenopodiales Chironiales Chlor

anthalesChrysobalanales Cinchonales Circaeasterales Cistales Citrales Cocosales Col

chicales Columelliales CombretalesCommelinales Connarales Convolvulales Coriarial

es Cornales Corylales Corynocarpales Crassulales CrossosomatalesCucurbitales Cunn

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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inghamiales Cunoniales Cupressales Cycadales Cyclanthales Cymodoceales Cynarales

CynomorialesCyperales Cyrillales Cytinales

Daphnales Daphniphyllales Dasypogonales Datiscales Degeneriales Desfontainiales D

iapensiales Didymelales DillenialesDioales Dioncophyllales Dioscoreales Diospyrales

Dipentodontales Dipsacales Droserales

Ebenales Echiales Elaeagnales Elaeocarpales Elatinales Elodeales Emmotales Empetr

ales Ephedrales Ericales EriocaulalesErythropalaless Escalloniales Eucommiales Euph

orbiales Eupomatiales Eupteleales Euryalaless

Fabales Fagales Falcatifoliales Ficales Flacourtiales Flagellariales Fouquieriales Franc

oales Frangulales

Galiales Garryales Geissolomatales Gentianales Geraniales Gesneriales Ginkgoales G

laucidiales Globulariales GnetalesGoodeniales Greyiales Griseliniales Grossulariales

Grubbiales Gunnerales Gyrocarpales Gyrostemonales

Haemodorales Haloragales Hamamelidales Hanguanales Heisteriales Helleborales He

lwingiales Himantandrales HippuridalesHomaliales Hortensiales Huales Huerteales

Hydatellales Hydnorales Hydrangeales Hydrastidales HydrocharitalesHydropeltidales

Hydrostachyales Hypericales Hypoxidales

Icacinales Illiciales Iridales Irvingales Iteales Ixiales

Jasminales Juglandales Julianiales Juncaginales Juncales

Lacistematales Lactoridales Lamiales Lardizabalales Laurales Lecythidales Ledocarpal

es Leitneriales LentibularialesLigustrales Liliales Limnanthales Linales Liriales Loasal

es Lobeliales Loganiales Lonicerales Loranthales LowialesLythrales

Magnoliales Malpighiales Malvales Marathrales Marcgraviales Mayacales Medusagy

nales Medusandrales MelanthialesMelastomatales Meliales Melianthales Meliosmal

es Menispermales Menyanthales Metteniusales MitrastemonalesMiyoshiales Moni

miales Moringales Musales Myricales Myristicales Myrothamnales Myrsinales Myrta

les

Najadales Nandinales Narcissales Nartheciales Nelumbonales Nepenthales Neuradal

es Nitrariales Nolanales NothofagalesNyctaginales Nymphaeales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Ochnales Oenotherales Olacales Oleales Onagrales Oncothecales Opuntiales Orchid

ales Oxalidales

Saxifragales Pandales Pandanales Papaverales Paracryphiales Parasitaxales Paridales

Parnassiales PassifloralesPenaeales Pennantiales Pentaphragmatales Peridiscales P

etiveriales Petrosaviales Phellinales Philydrales PhyllanthalesPhysenales Phytolaccal

es Picramniales Pinales Pinguiculales Piperales Pittosporales Plantaginales Platanale

sPlumbaginales Poales Podocarpales Podophyllales Podostemales Polemoniales Pol

ygalales Polygonales PontederialesPortulacales Posidoniales Potamogetonales Primu

lales Proteales

Quercales Quillajales Quintiniales

Rafflesiales Ranunculales Rapateales Resedales Restionales Rhabdodendrales Rhamn

ales Rhinanthales RhizophoralesRhodorales Rhoipteleales Roridulales Rosales Rouss

eales Rubiales Ruppiales Rutales

Sabiales Salicales Salvadorales Samolales Samydales Sanguisorbales Santalales Sapin

dales Sapotales SarracenialesSaxegotheales Saxifragales Scheuchzeriales Sciadopitya

les Scleranthales Scrophulariales Scyphostegiales SedalesSilenales Simmondsiales S

milacales Solanales Sphenocleales Sphenostemonales Stangeriales Stellariales Stemo

nalesStilbales Stylidiales Styracales Surianales

Taccales Tamales Tamaricales Taxales Taxodiales Tecophilaeales Terebinthales Tern

stroemiales Theales TheligonalesThymelaeales Tiliales Tofieldiales Torricelliales Tov

ariales Tribelales Trilliales Trimeniales Triuridales TrochodendralesTropaeolales Turn

erales Typhales

Ulmales Urticales

Vacciniales Vahliales Vallisneriales Velloziales Veratrales Verbenales Viburnales Vinc

ales Violales Viscales VitalesVochysiales

Welwitschiales Winterales

Xanthorrhoeales Ximeniales Xyridales

Zamiales Zingiberales Zosterales Zygophyllales

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which the genus of the basionym is to be found

Abietaceae Abolbodaceae Abrophyllaceae Acalyphaceae Acanthaceae [Acanthochla

mydaceae] Acarnaceaee AceraceaeAchariaceae Achatocarpaceae Achradaceae Achy

ranthaceae Acmopylaceae Acoraceae Acristaceae ActaeaceaeActinidiaceae Actinota

ceae Amentotaxaceae Adenogrammaceae Adoxaceae Aegialitidaceae Aegicerataceae

Aegilopaceae Aeginetiaceaee Aegiphilaceae Aesculaceae Aextoxicaceae Agapanthac

eae Agavaceae AgdestidaceaeAgialidaceae Agrimoniaceae Agrostidaceae Ailanthace

ae Aitoniaceae Aizoaceae Akaniaceae Alangiaceae AlchemillaceaeAldrovandaceae A

lismataceae Alliaceae Allioniaceae Allophylaceae Aloaceae Alopecuraceae Alpiniacea

e AlseuosmiaceaeAlsinaceae Alsodeiaceae Alstroemeriaceae Altingiaceae Alzateacea

e Amaranthaceae Amaryllidaceae Amborellaceae Ambrosiaceae Amentotaxaceae A

mmanniaceae Ammiaceae Amomaceae Ampelidopsaceae Amygdalaceae Amyridacea

eAnacardiaceae Anagallidaceae Anarthriaceae Anchusaceae Ancistrocladaceae Andr

omedaceae AndropogonaceaeAndrostachyaceae Androsynaceae Anemarrhenaceae

Anemonaceae Angelicaceae Anisophylleaceae AnnonaceaeAnomochloaceae Anopter

aceae Anrederaceae Anthemidaceae Anthericaceae Anthobolaceae Antidesmataceae

Antirrhinaceae Antoniaceae Aparinaceae Aphanopetalaceae Aphloiaceae Aphyllanth

aceae Apiaceae ApocynaceaeApodanthaceae Aponogetonaceae Aporusaceae Apose

ridaceae Apostasiaceae Aptandraceae AquifoliaceaeAquilariaceae Aquilegiaceae A

raceae Aragoaceae Araliaceae Aralidiaceaee Araucariaceae Arbutaceae Arceuthidace

aeArceuthobiaceae Arctostaphylaceae Arctotidaceae Ardisiaceae Arecaceae Argoph

yllaceae Arisaraceae AristolochiaceaeAristoteliaceae Arjonaceae Armeriaceae Artem

isiaceae Arthrotaxidaceae Artocarpaceae Arundinaceae ArundinellaceaeAsaraceae A

sclepiadaceae Ascyraceaee Asiraceae Asparagaceae Asperulaceae Asphodelaceae As

pidistraceaeeAsteliaceae Asteraceae Asteranthaceae Asteropeiaceae Astragalaceae

Astrocarpaceae AthanasiaceaeAtherospermataceae Atriplicaceae Atropaceae Aucuba

ceae Aurantiaceae Austrobaileyaceae Austrotaxaceae AvenaceaeAverrhoaceae Avet

raceae Avicenniaceae Azaleaceae Azimaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Balanitaceae Balanopaceae Balanophoraceae Balsameaceae Balsaminaceae Bambusa

ceae Barbaceniaceae BarbeuiaceaeBarbeyaceae Barclayaceae Barringtoniaceae Base

llaceae Bataceae Baueraceae Bauhiniaceae Baxteriaceae BegoniaceaeBehniaceae B

elangeraceae Belloniaceae Bembiciaceae Berberidaceae Berberidopsidaceae Berryac

eaee BersamaceaeBertyaceae Berzeliaceae Besleriaceae Betaceae Betulaceae Biebe

rsteiniaceae Bifariaceae Bignoniaceae BischofiaceaeBixaceae Blakeaceae Blandfordia

ceae Blattiaceae Blepharocaryaceae Blitaceae Blyxaceae Boerlagellaceae Bolivaracea

eBombacaceae Bonnetiaceae Bontiaceae Boopidaceae Boraginaceae Borassaceae B

oroniaceae BoryaceaeBotryodendraceae Bougainvilleaceae Boweniaceae Brassicacea

e Brachycaulaceae Bretschneideraceae BrexiaceaeBromeliaceae Brownlowiaceae Br

unelliaceae Bruniaceae Brunoniaceae Brunsvigiaceae Bryoniaceae BuchneraceaeBuci

daceae Buddlejaceae Buglossaceae Bulbocodiaceae Bumeliaceae Bupleuraceae Burc

hardiaceae BurmanniaceaeBurseraceae Butneriaceae Butomaceae Buxaceae Byblida

ceae Byttneriaceae

Cabombaceae Cacaoaceae Cactaceae Caesalpiniaceae Caladiaceae Calamaceae Calce

olariaceae CalectasiaceaeCalendulaceae Callaceae Callicomaceae Calligonaceae Calli

traceae Callitrichaceae Calochortaceae CalophyllaceaeCalthaceae Calycanthaceae Ca

lyceraceae Cambogiaceae Camelliaceae Campanulaceae CampynemataceaeCanacom

yricaceae Canellaceae Cannabaceae Cannaceae Canopodaceae Canotiaceae Cansjera

ceae CapparaceaeCaprariaceae Caprifoliaceae Cardiopteridaceae Carduaceae Carica

ceae Carissaceae Carlemanniaceae CarpinaceaeCarpodetaceae Cartonemataceae Ca

ryocaraceae Caryophyllaceae Casearieaceae Cassiaceae CassipoureaceaeCassythacea

e Castaneaceae Castelaceae Casuarinaceae Catesbaeaceae Cathedraceae Cecropiace

ae CedraceaeCedrelaceae Celastraceae Celosiaceae Celtidaceae Centaureaceae Cen

trolepidaceae Centroplacaceae CepaceaeCephalanthaceae Cephalotaceae Cephalota

xaceae Cerastiaceae Ceratoniaceae Ceratophyllaceae CerberaceaeCercidiphyllaceae

Cercocarpaceae Cercodiaceae Cereaceae Cerinthaceae Ceroxylaceae Cestraceae Cev

alliaceaeChailletiaceae Chamaedoraceae Chamaemoraceae Chamelauciaceae Chaun

ochitonaceae Chelidoniaceae ChelonaceaeChenopodiaceae Chimonanthaceae Chingi

thamnaceae Chionographidaceae Chiranthodendraceae ChironiaceaeChlaenaceae Ch

loanthaceae Chloranthaceae Chloridaceae Chlorogalaceae Chrysobalanaceae Cicerac

eae CichoriaceaeCimicifugaceae Cinchoniaceae Circaeaceae Circaeasteraceae Cissac

eae Cistaceae Citraceae Clematidaceae CleomaceaeClethraceae Clusiaceae Cneorac

eae Cnestidaceae Cnicaceae Cobaeaceae Cochlospermaceae Cocosaceae Coffeaceae

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Colchicaceae Coleogynaceae Columelliaceae Combretaceae Commelinaceae Comocl

adiaceae Compositae CompsoaceaeConantheraceae Connaraceae Conostylidaceae C

onvallariaceae Convolvulaceae Cordiaceae CoreopsidaceaeCoriandraceae Coriariacea

e Coridaceae Corispermaceae Cornaceae Corokiaceae Coronillaceae Correaceae Cor

rigiolaceaeCorsiaceae Corydalaceae Corylaceae Corynocarpaceae Coryphaceae Cost

aceae Cotyledonaceae CoulaceaeCoutariaceae Coutoubiaceae Crassulaceae Crescen

tiaceae Cressaceae Crinaceae Crocaceae CroomiaceaeCrossosomataceae Crotonacea

e Cruciferae Cryptaceae Crypteroniaceae Cryptocorynaceae CryptomeriaceaeCtenol

ophonaceae Cucurbitaceae Cunninghamiaceae Cunoniaceae Cupressaceae Curcumac

eae Curtisiaceae CuscutaceaeCyananthaceae Cyanastraceae Cynanchaceae Cyanella

ceae Cycadaceae Cyclanthaceae CyclantheraceaeCyclocheilaceae Cydoniaceae Cymo

doceaceae Cynaraceae Cynocrambaceae Cynomoriaceae Cyperaceae CyphiaceaeCyp

hocarpaceae Cypripediaceae Cyrillaceae Cyrtandraceae Cyrtanthaceae Cytinaceae

Dacrycarpaceae Dactylanthaceae Damasoniaceae Daphnaceae Daphniphyllaceae Das

ypogonaceae DatiscaceaeDaturaceae Daucaceae Davidiaceae Davidsoniaceae Decais

neaceae Deeringiaceae Degeneriaceae DelphiniaceaeDendrophthoaceae Columelliac

eae Detariaceae Dialypetalanthaceae Dianellaceae Dianthaceae DiapensiaceaeDicha

petalaceae Dichondraceae Diclidantheraceae Dicrastylidiaceae Dictamnaceae Didiere

aceae DidymelaceaeDidymocarpaceae Diegodendraceae Diervillaceae Digitalidaceae

e Dilatridaceae Dilleniaceae Dioaceae DionaeaceaeDioncophyllaceae Dioscoreaceae

Diosmaceae Diospyraceae Dipentodontaceae Diphylleiaceae Diplolaenaceae Dipsaca

ceaeDipterocarpaceae Dirachmaceae Disanthaceae Diselmaceae Dodonaeaceae Do

mbeyaceae Donatiaceae DorsteniaceaeDortmannaceae Doryanthaceae Drabaceae D

racaenaceae Dracontiaceae Drimyidaceae Droseraceae DrosophyllaceaeDryadaceae

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aceae

Ebenaceae Eccremidaceae Ecdeiocoleaceae Echiaceae Echinopaceae Ehretiaceae Ela

eagnaceae ElaeocarpaceaeElegiaceae Elatinaceae Ellisiaceae Ellisiophyllaceae Elodea

ceae Elytranthaceae Embeliaceae Emblingiaceae EmmotaceaeEmpetraceae Encepha

lartaceae Engelhardtiaceae Enhalaceae Epacridaceae Ephedraceae Ephemeranthacea

eEpilobiaceae Epimediaceae Eragrostidaceae Eremolepidaceae Eremosynaceae Erica

ceae Erinaceae EriocaulaceaeEriogonaceae Eriospermaceae Erodiaceae Erycibaceae

Eryngiaceae Erysimaceae Erythroniaceae ErythropalaceaeErythrospermaceae Erythro

xylaceae Escalloniaceae Eschscholziaceae Eucomidaceae Eucommiaceae Eucryphiace

aeEuonymaceae Eupatoriaceae Euphorbiaceae Euphrasiaceae Euphroniaceae Eupo

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matiaceae Eupteleaceae EuryalaceaeEustrephaceae Euthemidaceae Eutocaceae Exb

ucklandiaceae Exocarpaceae

Fabaceae Fagaceae Falcatifoliaceae Ferulaceae Festucaceae Ficaceae Ficoidaceae Fit

zroyaceae FlacourtiaceaeFlagellariaceae Flindersiaceae Foetidiaceae Forestieraceae

Fothergillaceae Fouquieriaceae Fragariaceae FrancoaceaeFrangulaceae Frankeniacea

e Fraxinaceae Fraxinellaceae Freycinetiaceae Fritillariaceae Fuchsiaceae Fumariacea

eFunkiaceae

Gaiadendraceae Galacaceae Galanthaceae Galaxiaceae Galedupaceae Galeniaceae G

aliaceae Garciniaceae GardeniaceaeGardneriaceae Garryaceae Geissolomataceae Ge

itonoplesiaceae Gelsemiaceae Geniostomaceae GentianaceaeGeonomataceae Geosir

idaceae Geraniaceae Gerrardinaceae Gesneriaceae Gethyllidaceae Gilliesiaceae Gina

lloaceaeGinkgoaceae Gisekiaceae Gladiolaceae Glaucidiaceae Glechomaceae Glinace

ae Globulariaceae Gnaphaliaceae GnetaceaeGoetzeaceae Gomortegaceae Gomphiac

eae Gomphrenaceae Gonystylaceae Goodeniaceae Gordoniaceae GouaniaceaeGoup

iaceae Gramineae Gratiolaceae Grewiaceae Greyiaceae Grielaceae Griseliniaceae Gr

onoviaceae GrossulariaceaeGrubbiaceae Guaiacanaceae Guamatelaceae Guettardac

eae Gunneraceae Gustaviaceae Guttiferae GyrocarpaceaeGyrostemonaceae

Hachetteaceae Haemanthaceae Haemodoraceae Halesiaceae Hallieraceae Halocarpa

ceae Halophilaceae HalophytaceaeHaloragaceae Hamamelidaceae Hanguanaceae H

aptanthaceae Harmandaceae Hebenstretiaceae HectorellaceaeHederaceae Hedyosm

aceae Hedyotidaceae Hedysaraceae Heisteriaceae Heleniaceae Heliamphoraceae H

elianthaceaeHelianthemaceae Helichrysaceae Heliconiaceae Helicteraceae Heliotropi

aceae Helleboraceae Heloniadaceae HelosidaceaeHelwingiaceae Hemerocallidaceae

Hemimeridaceae Hemiphylacaceae Henriqueziaceae Henslowiaceae Hermanniaceae

Hernandiaceae Herniariaceae Herreriaceae Hesperocallidaceae Heterantheraceae He

teropyxidaceae HeterostylaceaeHibbertiaceae Hibiscaceae Hilleriaceae Himantandra

ceae Hippocastanaceae Hippocrateaceae HippomanaceaeHippophaeaceae Hippurida

ceae Hirtellaceae Holacanthaceae Homaliaceae Hopkinsiaceae Hoplestigmataceae H

ordeaceaeHornschuchiaceae Hortensiaceae Hortoniaceae Hostaceae Houstoniaceae

Huaceae Huerteaceae HugoniaceaeHumbertiaceae Humiriaceae Hyacinthaceae Hyd

atellaceae Hydnoraceae Hydrangeaceae Hydrastidaceae HydrillaceaeHydrocharitacea

e Hydrocotylaceae Hydrogetonaceae Hydroleaceae Hydropeltidaceae Hydrophylacac

eae HydrophyllaceaeHydrostachyaceae Hymenocardiaceae Hypecoaceae Hyperanth

eraceae Hypericaceae Hypopityaceae HypoxidaceaeHypseocharitaceae

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Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta

ceae ImperatoriaceaeInocarpaceae Inulaceae Ionidiaceae Iriarteaceae Iridaceae Irvi

ngiaceae Isnardiaceae Isophysidaceae IteaceaeStrasburgeriaceae Ixiaceae Ixioliriace

ae Ixonanthaceae

Jalapaceae Jamboliferaceae Japonoliriaceae Jasionaceae Jasminiaceae Johnsoniaceae

Joinvilleaceae JuglandaceaeJulianiaceae Juncaceae Juncaginaceae Juniperaceae Jus

siaeaceae Justiciaceae

Kadsuraceae Kaliphoraceae Kalmiaceae Kaniaceae Kiggelariaceae Kingdoniaceae Kin

giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae

Krameriaceae

Labiatae Lacandoniaceae Lachenaliaceae Lacistemataceae Lactoridaceae Lactucaceae

Lagerstroemiaceaee LamiaceaeLampsanaceae Lanariaceae Langsdorffiaceae Lantan

aceae Lapageriaceae Lardizabalaceae Lasiopetalaceae LauraceaeLawsoniaceae Laxm

anniaceae Lecythidaceae Ledaceae Ledocarpaceae Leeaceae Leguminosae Leitneriac

eae LemnaceaeLennoaceae Lentibulariaceae Lentiscaceae Leoniaceae Leonticaceae

Lepidobotryaceae LepidocarpaceaeLepidocaryaceae Lepidocerataceae Leptaulaceae

Leptospermaceae Lepturaceae Lepuropetalaceae LeuctenbergiaceaeLibocedraceae

Licaniaceae Ligustraceae Lilaeaceae Liliaceae Limeaceae Limnanthaceae Limnocharit

aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace

ae Linderniaceae Lindleyaceae Linnaeaceae LippayaceaeLiriaceae Liriodendraceae Li

ssocarpaceae Littorellaceae Loasaceae Lobeliaceaee Loganiaceae LomandraceaeLoni

ceraceae Lopeziaceae Lophiocarpaceae Lophiolaceae Lophiraceae Lophophytaceae L

ophopyxidaceae LoranthaceaeLotaceae Lowiaceae Lupulaceaee Luxemburgiaceae Lu

zuriagaceae Lychnidaceae Lyciaceae LyginiaceaeLygodisodeaceae Lysimachiaceae Ly

thraceae

Macarisiaceae Mackinlayaceae Madiaceae Maesaceae Magnoliaceae Malaceae Male

sherbiaceae MalortieaceaeMalpighiaceae Malvaceae Mangiaceae Manicariaceae Ma

paniaceae Marantaceae Marathraceae MarcgraviaceaeMartyniaceae Mastixiaceae

Matricariaceae Maundiaceae Mayacaceae Medeolaceae Medusagynaceae Peridiscac

eaeMelaleucaceae Melampyraceae Melanophyllaceae Melanthiaceae Melastomatac

eae Meliaceae Melianthaceae MelicaceaeMeliosmaceae Melittidaceae Melochiacea

e Memecylaceae Mendonciaceae Menispermaceae Menthaceae MenyanthaceaeMe

nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae

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Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

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Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

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ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

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hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

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Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

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heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

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Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

49

cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

50

ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 16: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

16

content of the clades has changed considerably and incidentally they are now easier to

identify than before the decision to recognise two families is not difficult The same is

true for Salicaceae and Achariaceae (Malpighiales) two previously small families that

have received the bulk of the old Flacourtiaceae Clade and hence taxon limits remain

difficult around Theaceae (Ericalessee alsoSladeniaceae Pentaphylacaceae)

and Euphorbiaceae (Malpighiales see

also Phyllanthaceae Picrodendraceae Putranjivaceae etc) Here current groupings

may not represent quite such dramatic changes in our understanding of relationships

Indeed parts of the old Euphorbiaceae that were separated may yet go back together

particularly Phyllanthaceae and Picrodendraceae (see Wurdack et al 2004 Davis et al

2005) even if Theaceae should indeed be dismembered (Geuten et al 2004) However

given that there is no molecular evidence that currently warrants combining all the

segregates of Euphorbiaceae and even if some do come together the clades the

families represent suggest novel groupings not recognised in current classifications

Furthermore Rafflesiaceae appear to be embedded within Euphorbiaceae s str (Davis

et al 2007) so the family is divided Of course maintaining even a moderately narrowly-

circumscribed Euphorbiaceae would entail reducing the iconic Rafflesiaceae to

synonymy Relationships in core Caryophyllales especially

around Phytolaccaceae andNyctaginaceae and also Portulacaceae are incompletely

understood and refashioning of taxon limits will doubtless be needed as cladistic

relationships become apparent (eg see Nyffeler 2007 Ogburn 2007a b) Some

groupings in the old Icacinaceaeand Olacaceae also remain unclear

The discovery of the relationships of parasitic and aquatic groups have presented a

particular challenge to systematists Morphologically some of these plants are so highly

modified that interpretation of the plant body in conventional terms is difficult or even

impossible Thus parasitic groups (broad-leaved angiosperms only) such as Rafflesiaceae

are hard to place since both the vegetative body and the flowers are changed almost

beyond recognition (flowers of rafflesiaceae are a good example of gigantism - Davis et

al 2007) Furthermore plastid gene sequences may be difficult or impossible to obtain

the chloroplast DNA in particular being highly degraded and the rate of molecular

change in general may be high so the problem of long-branch attraction is serious (eg

Duff amp Nickrent 1997 Nickrent et al 1998 Caddick et al 2002a G Petersen et al

2006b) Echlorophyllous sapromycoheterotrophic taxa mostly known from monocots

present similar problems However progress is being made placements for

Apodanthaceae Rafflesiaceae Mitrastemonaceae Cytinaceae and Cynomoriaceae have

recently been suggested (Barkman et al 2004 2007 Davis amp Wurdack 2004 Nickrent et

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

17

al 2004 Davis et al 2007) it seems likely that Burmanniaceae sl are polyphyletic but

both parts are in Dioscoreales (Merckx et al 2006) and relationships within the largely

hemiparasitic Santalales are also gradually being clarified (Maleacutecot 2002) The inclusion

of parasitic taxa in general molecular analyses can cause conniptions (eg Nickrent et al

2004 Davis et al 2004 Chase et al 2006 G Petersen et al 2006b) and there can be

horizontal transmission of genes (eg Davis amp Wurdack 2005 Vitaceae to Rafflesiaceae

Barkman et al 2007 the mitochondrial atp1 gene commonly moves)

Similarly in water plants neither vessels in particular nor much xylem in general is

needed leaves are highly modified and water-mediated pollination if adopted may

well be associated with major changes in floral morphology Here too recent molecular

studies suggest that aquatic groups with hitherto problematic relationships may find

homes Thus Podostemaceae are close to Clusiaceae and Calophyllaceae and are sister

to Hypericaceae (Malpighiales Kita amp Kato 2001) Hydatellaceae which used to be in

Poales are part of Nymphaeales (Saarela et al 2007) and Hydrostachyaceae may be

close to Hydrangeaceae (Cornales Xiang et al 2002) or perhaps should be placed in

Lamiales note that in the first two cases in particular there are morphological and

chemical features that support such a move If Podostemaceae are indeed close to

Clusiaceae I look forward to seeing hypotheses to explain how the dramatic changes in

the vegetative body that have made Podostemaceae so problematic for generations of

systematists took place That conventional wisdom has trouble in understanding or

explaining how the morphologies of groups like Clusiaceae and Podostemaceae can be

related is largely a problem with conventional wisdom Ceratophyllaceae seem to be

finding a position as sister to eudicots (see Moore et al 2007) but their morphology is

so derived that there is no morphological evidence of which I am aware for this

relationship There are also similar although less extreme morphological problems with

many plants pollinated by wind

In such situations especially with some water plants mycoheterotrophs and

parasites a variety of characters may be affected and particularly under such

circumstances groups have tended to be formed using a few or even only a single

character that seems to provide evidence of relationships Add to this the tendency to

weight some characters particularly strongly on a priori grounds and the result was

either the recognition of conglomerate taxa such as Amentiferae which now appear to

be highly polyphyletic or the segregation of families like Plantaginaceae s str (now

much expanded) and Leitneriaceae (now in Simaroubaceae) - all of these are more or

less wind pollinated

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

18

Even although relationships of many parasitic and aquatic groups are being

clarified and at least some of the proposed phylogenetic positions seem to be well

supported it is indeed often still very difficult to understand how such groups have

evolved from their more morphologically conventional relatives However we have had

all sorts of preconceptions as to how characters should evolve Thus we used to

assume that features like highly scalariform vessel perforation plates or the complete

absence of vessels or a flower with an androecium that had many stamens a superior

ovary or separate petals were necessarily plesiomorphic or primitive and conversely

a simple perforation plate and the presence of vessels an androecium with few

stamens inferior ovary or petals that were connnate were almost necessarily

apomorphic or advanced (but cf eg Stebbins 1951) Such assumptions are incorrect

(eg Soltis et al 2005b) Carpels may become secondarily free carpels may fail to close

the seeds then developing outside the confines of the carpel as in some Aspagaraceae-

Nolinoideae Violaceae Berberidaceae Malvaceae-Sterculioideae etc) in Peliosanthes

teta perhaps the only species in Peliosanthes(Asparagaceae-Nolinoideae) the ovary

varies from superior to inferior (Jessop 1976 species limits here need close

investigation see also Kuzoff et al 2001 and Soltis amp Hufford 2002 Saxifragaceae

Apiales etc) many-seeded carpels can evolve from few-seeded carpels

(Razafimandimbison et al 2008) monoecy may be derived from dioecy (Schaefer amp

Renner 2010 and references) Classic studies such as those by Babcock (eg 1947)

on Crepis that assumed that evolution - in this case of the karyotype in particular - was

unidirectional have needed comprehensive re-evaluation (Enke amp Gemeinholzer 2008)

Most if not all characters have reversed andor evolved in parallel as is clear in the

discussion of ovary position here especially in Poales Saxifragales and Asparagales and

also in comments on such morphologically distinctive taxa (when compared with their

immediate relatives) such as Menyanthaceae and especially Pittosporaceae Parallel

evolution may occur even at the level of amino acid substitution as in the independent

acquisition of the phosphoenolpyruvate carboxylase (pepC) gene in C4 photosynthesis in

grasses (Christin et al 2007b see also Blaumlsing et al 2000)

Thinking of how such characters have been used in the past in classifications at the

generic level the results have been similar to those just discussed above for parasitic

plants and others Relying too much on animal pollination syndromes to mark generic

boundaries has all too often led to taxa that are highly unsatisfactory phylogenetically

(see eg Acanthaceae Bignoniaceae Campanulaceae Ericaceae Lamiaceae

Orchidaceae) and over-reliance on characters of fruit and seed (see eg Brassicaceae

and Apiaceae) has also led to unsatisfactory generic limits Again the more general

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

19

problem is the use of one or a very few characters that have been weighted a priori to

structure classifications (see also Garciacutea et al 2009) although it is perhaps not

surprising that such an approach sometimes results in the delineation of taxa that have

indeed turned out to be quite distinct (cf eg Burtt Astragalus versus Oxytropis)

Taxa recognized here are monophyletic yet many are polythetic at the

morphological level that is they lack unique features characterizing (defining) all and

only members of that family they can be recognised phenetically only by the unique

combinations of characters that they posess This is the result of evolution any

synapomorphy characterizing a taxon may be lost or modified beyond easy recognition

in some of its members or the synapomorphy may appear to be identical to a feature

that has evolved in parallel in a quite unrelated plant That plant groups are polythetic is

almost as much a feature of monophyletic taxa as the paraphyletic taxa common in

evolutionary classifications Some families now include substantial variation as

phenetically distinct derived groups are placed in their proper phylogenetic position -

examples are Ericaceae which include the erstwhile Empetraceae a wind-pollinated

group that is florally very different and the overwhelmingly large-flowered and animal-

pollinated Plantaginaceae which now include several derived small-flowered aquatic

and wind-pollinated groups that were previously placed in separate families

Thus some families as delimited here may not be easy to recognize However

remember that detecting relationships - use whatever characters you can even if they

are not obvious - and naming a plant - focus on easy-to-see characters that may not

reflect relationships - are quite different problems Taxa although natural may not be

readily recognizable indeed it was in exactly this context that Lamarck worked out the

basic principles of writing dichotomous keys in 1778 (see Scharf 2007 for keys and the

like) Of course Lamarcks idea of nature was very different from ours - he thought that

there was some kind of continuum of form on which living organisms were to be

situated with no real gaps anywhere - but this meant that his genera (for example)

might well not be sharply distinct from each other even if each were part of the real

continuum that was life Lamarcks separation of the establishment of relationships and

groupings in a system and identification of the plants in that system have been widely

accepted (but cf eg Godfray amp Knapp 2004 many of the contributors to Heywood et

al 2007) Perhaps the best way of identifying plants at the family level is by well-made

multiple access keys as in Watson and Dallwitz (1992a onwards family limits there may

differ substantially from those adopted here) Multiple access keys free users from the

constraints of dichotomous keys in which particular characters are needed at each step

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

20

of the identification process before there can be further progress Instead those

characters that happen to be evident on a specimen can be used in whatever order is

convenient when linked to illustrations glossaries etc their power is enormous (see

Dallwitz et al 2000 [2006] for the principles underlying their construction and use)

Nevertheless dichotomous keys such as those of Hutchinson (1973) and Franz Thonner

(Geesink et al 1981) have their uses Of course taxa coming out adjacent in keys may

well not be at all related

(When identifying large numbers of plants even more efficient than either style of

identification and certainly lots more fun is sight identification Unless you have a

photographic memory you have to build up your knowledge of comparative plant

morphology - on which the ability to make accurate identifications depends - by

repeated observation When faced with an unknown plant I always look for leaf teeth

and stipules distinctive hair types especially stellate and T-shaped hairs glands and

punctations of any sort the presence of latex or other exudate and check leaf insertion

smelling crushed leaves can also be helpful In this context nodal anatomy can usually

be checked using a razor and a hand lens or even simply carefully examining leaf scars

The short paragraphs added after most families may help in confirming familial

identifications)

For the record and for the little that it is worth there are 4 orders and 13 families

of gymnosperms characterised on these pages and together they include some 82

genera and 947 species For angiosperms comparable figures are 56 orders 445

families 13208 genera and 261750 species (of which monocots include 11 orders 89

families 2759 genera and 52760 species) Note however that higher mathematics

was never my strong point and anyway these are pretty meaningless figures even for

species which many (but not all) might concede smacked slightly more of reality than

other taxa estimates range as high as 422000 (Govaerts 2001) Furthermore numbers

of genera and species change daily Nevertheless as emphasized here families are

useful in teaching we as a community can ensure that their limits remain largely stable

and by concentrating on relatively few of them one can gain some familiarity with much

of the worlds flora For summaries of the sizes of orders and families and of the general

arrangement followed in these pages see also the Statistics page itself

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

21

A PHYLOGENETIC TREE OF SEED PLANTS (THE MAIN TREE)

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

22

The Main Tree above is a conservative summary of well supported relationships

between the major clades of extant seed plants Reading - interpreting and

understanding - such a phylogenetic tree is not necessarily easy so reading a tree may

require a bit of practice OHara (eg 1988 1992) provides an accessible introduction to

thinking and talking about history in the context of looking at phylogenetic trees More

recently Krell and Cranston (2004) Crisp and Cook (2005) and others have emphasized

how careful one must be when interpreting and talking about evolution in ladderized

trees in particular and phylogenetic trees in general Gregory (2008) provides a

particularly detailed discussion of this problem In ladderized trees the smaller (in terms

of numbers of terminals) sister taxon is consistently shown on the same side at every

node thus the trees tend to be pectinate like the teeth of a comb however the

horizontal axis that is as a result so evident has no polarity or particular direction The

use of the adjective basal when discussing phylogenetic trees is especially dangerous

(see also Wojciechowski et al 2004 D Soltis et al 2005b) When I use the term and the

context is not otherwise clear I am referring to the pectinations at the base of a

ladderised tree Remember that when talking about sister taxa one can never be basal

to the other or older or younger although members of one can be more derived - in the

sense of having more apomorphies - than the other (but even then one has to be

careful) The words primitive and advanced and lower and higher should

practically never be used especially when talking about taxa Amborellaceae and

Pinaceae are sister to all other flowering plants and Pinales and are basal to the crown

groups of all other flowering plantsPinales respectively but that does not mean they

are thereby primitive Pinaceae in particular have numerous apomorphies The word

plesiomorphic is far less loaded than primitive and can be used to talk about

individual characters

Most trees here have been more or less ladderized as is the one above The

ladderization may be imperfect for example you can see that on the Main Tree

asterids with ten orders follow rosids and relatives which have seventeen although

this is in part because of the currently unresolved nature of relationships between the

main clades of core eudicots (this is likely to change soon) The Main Tree could have

been drawn with Amborella Acorus or a host of other taxa at the far right without

offending any relationships phylogenetic trees are like mobiles the only fixed points

being the nodes Despite the problems just mentioned pectination interpreted

carefully has its value and it is not an arbitrary process As one reads the terminals of a

pectinate tree from left to right adjacent terminals will be separated by apomorphies

that will cumulate in an appropriate fashion When thinking of a book or a herbarium

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

23

sequence (see Haston et al 2007) this is of some value Nymphaeales and

Austrobaileyales are here adjacent on the tree but they could be separated by

hundreds of families in the sequence merely by rotating only the node from which

Nymphaeales and their sister group (all other angiosperms minus about 75 species)

arise If adjacent in a book or herbarium or as in these summary pages then it is

relatively easy to relate their characters both apomorphic and plesiomorphic but if

separated by hundreds of pages or two floors in a large building then it is less easy to

get anything from the sequence (Since all orders in the pages of the main part of this

site are preceded by the apomorphies of all nodes immediately below them in the seed

plant phylogeny and because of the linkages that have been built in to the site this

problem is much less serious there there is no sequence) So the particular sequence of

all seed plants that is based on a phylogenetic classification but is to be used in herbaria

or any other place where taxa have to be arranged linearly as in this part of APweb

will be one that maximizes the number of taxa that are both successive branches of the

tree and placed successively in the sequence Since specimens are generally filed under

families the outline of a new family sequence for arranging herbaria and books can now

be suggested (Haston et al 2007) and such a standardized sequence will further help

teaching and learning about plants

ON THINKING ABOUT APOMORPHIES

Identifying apomorphies is important because understanding the evolution of

morphology in the broad sense - ie including anatomy chemistry etc - in the context

of diversification is one of our major goals For identifying apomorphies several

preconditions must be met One needs to have an accurate well supported phylogeny

one has to have examined the right taxa both from the point of view of morphology and

molecules one has to have coded the characters right (ie delimited states

appropriately) and one has to use an appropriate model of evolution when fitting the

variation to the tree (see eg Omland 1999 Stevens 2006b) If the distributions of

apomorphies below are compared with those in other studies (eg Turgeon et al 2001

Bremer et al 2001 Endress 2001 Albach et al 2001a Judd amp Olmstead 2004 D Soltis

et al 2005b Zhang et al 2006 Judd et al 2007) the reader may find a number of

differences Although I have integrated such studies as far as is possible there are five

reasons that link to the preconditions just mentioned why there may be differences

where features are placed on trees

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

24

1 I may not have found all the information about a particular character there may be

disagreement over its interpretation or I have added information

2 The sampling of nearly all molecular studies is very incomplete (see Salisbury amp Kim

2001 for problems caused by sampling) indeed it is commonplace to decry the

incompleteness of molecular sampling But not only is the sampling in molecular studies

often less than we might wish that of the morphological and chemical characters whose

evolution we are interested in understanding is also often very poor So for many

anatomical chemical and embryological characters that are confidently said to

characterise families and other groups we all too often have no idea if those characters

are applicable to the whole clade or just to a subgroup within that clade However as

our sampling improves we can locate changes on the tree more precisely if in

unexpected positions Thus is is sometimes suggested that Ericaceae have ellagic acid

(Soltis et al 2005b) in fact the little evidence we have indicates that only a small clade

of some 80 species has ellagic acid and the rest of the family - itself alone about one

third of all Ericales - does not Similarly Ericaceae are noted for their

ectendomycorrhizal associations but Enkianthus sister to all other Ericaceae

apparently lacks these (Abe 2005) Thus these mycorrhizae and their associated

distinctive hair roots are unlikely to be apomorphies of Ericaceae - however it appears

that other families near Ericaceae may also have ectendomycorrhizae (see Asai 1934)

3 I am fitting characters to a very conservative tree with many polytomies although the

nodes that are utilised are for the most part strongly supported polytomies make the

optimisation of characters that is the assigment of character state change to a

particular node on the tree notably difficult (eg Madison amp Madison 2002) In nearly

all studies of the evolution of characters distributions of characters are optimised on a

more or less fully resolved tree and the construction of supertrees may yield yet more

detailed hypotheses of relationships (for literature on supertrees see Cotton amp

Wilkinson 2007 2008) Of course some nodes on such fully resolved trees andor

supertrees may have little support and optimisations of characters on such trees may

carry correspondingly little conviction Even parts of some of the trees used here have

poor support eg relationships within aquatic Alismatales etc although I indicate

when this is the case

4 Exactly how one goes about optimising a character on a tree is critically important

Even using simple parsimony optimisations (ACCTRAN or DELTRAN ACCelerated

TRANsitions or DELayed TRANsitions) the position of synapomorphies on trees - and

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

25

hence our ideas of evolution - may differ and this is still more true if one uses maximum

likelihood or Bayesian methods and makes apparently reasonable suggestions about

weighting gains over losses (eg Donoghue amp Ackerley 1996 Cunningham et al 1998

Omland 1997 1999 Ree amp Donoghue 1999 Polly 2001 Webster amp Purvis 2001

Ronquist 2004 Crisp amp Cook 2005) Sannier et al (2007) show how in in Arecaceae that

where on a phylogenetic tree one might peg changes in microsporogenesis will depend

on the methods one uses to do this and Pedersen et al (2007) discuss the sometimes

very substantial effect of node support on the posterior probabilities of ancestral

character states Here I use parsimony optimization not always as explicit as it might be

but I have often indicated where there are particularly important uncertainties as to the

positions of particular character changes on the tree

5 Finally although I have paid quite a lot of attention to the delimitation of the

character states that make up all the characterizations I have not spent enough time on

this critical operation If we are interested in understanding evolution then fitting the

basic variation - not character states - to a tree in principle allows greater flexibility in

understanding morphology in the context of local phylogenies (see also Stevens 2000

Endress 2005c) However many character states used here are delimited globally that

is they are circumscribed in the context of the variation shown by individual characters

across all angiosperms andor in the context of classic ideas of character evolution

Character states often have arbitrary limits and serve best to communicate

information whether they are in fact suitable for either phylogenetic analysis or

understanding evolution are separate issues Studies have rather unsurprisingly

perhaps but importantly shown that dividing the one character into different sets of

states may yield differing ideas of evolution of that character (eg Lamb Frye amp Kron

2003 Hibbett 2004) When looking at trees on which character states are optimised

one should bear in mind the problems surrounding the delimitation of states (eg

Stevens 2000 2006b) and the danger of using pollen or other types - constructs

based on many characters that vary independently but which effectively get lost in

these types thus Blackmore et al (2009) decomposed the pollen types in Asteraceae

into 52 characters

I use Remanes three main criteria of homology or better similarity when

determining the basic similarity of structures on different organisms (see Remane 1952)

These criteria are special properties position and intermediates Special properties

include anatomical or chemical characters gene expression data etc Position refers

to the position of an organ with respect to landmarks on the plant Although plants are

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

26

plants and landmarks are not as fixed as one might like trying to work out the relative

positions of parts is a good way of understanding morphology so long as one does not

try to out-do Procrustes Intermediates include intermediates found on the same

plant or on different plants Intermediates may observed during development or be

adult structures Thus one may find changes during development which suggest what

the nature of a particular structure is and very different stuctures on different plants

may turn out to be similar early in development Or one can simply compare different

taxa and see that structures that appear to be very different when comparing two

different taxa can be linked morphologically by looking at other taxa

Nevertheless the use of these criteria may not yield an unambiguous answer as to

what a structure is even given a solid phylogeny and an improved understanding of

development (see Jaramillo amp Kramer 2007 for a useful discussion) As Endress (2005c)

observed a number of features - position function development shape anatomy

histology gene activity and relationships to other taxa that clearly have petals - can be

used to distinguish a petal (for example) from other floral structures if a petal does not

have one of these features is it thereby not a petal Thus Maturen et al (2005) recently

found that floral organ diversity genes (B and C) were expressed in the large white

inflorescence bracts of Cornus (see also Costa et al 2005) Peney et al (2005) noted that

not all monosulcate pollen grains in monocots have the same developmental pathway

and that as a result such pollen might not have the same ancestral state Reeves and

Olmstead (2003) suggested that the genetic mechanisms causing monosymmetry in

Lamiales and Solanales were different and Serna and Martin (2006) described similar

problems with the development of hairs in Arabidopsis when compared with that of

hairs in Antirrhinum and Solanaceae Indeed as one perhaps might expect delimitation

of states and characters does not necessarily become easier with increasing knowledge

of development etc Thus Buzgo et al (2004) Matthews and Endress (2005) and others

have shown how hard it can be to distinguish between eg prophylls and other floral

structures as their behaviour is studied during the course of the development of the

flower Similarly at what concentration is a particular secondary metabolite deemed to

be present (Waterman (2007)

The validity of the approach used here that of fitting morphological variation to a

largely molecular-based tree may be questioned However I think it rather unlikely that

well-supported molecular branches will be overturned by morphological data Indeed

analyses of morphological data alone do provide support for many of the clades evident

in molecule-only analyses and analyses with morphological and molecular data

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

27

together may lead to increased support for clades (eg Hufford 1992 Nandi et al 1998

[but adding morphological data reduces support for a number of critical clades too]

Doyle amp Endress 2000 however in none of these papers is the use of morphology

without ambiguity) It is unfortunately clear that the use of morphology alone may not

suggest problems in the phylogenetic placement of taxa that later turn out to have been

wrongly included (eg Zhang et al 1992) Thus I have been wary of putting much weight

on clades that have only morphological support but note that I have treated molecule-

based clades with low bootstrap or jacknife support values (esp below 70) or low

posterior probabilies (below 095) likewise Although I may have been mistaken in

placing so much emphasis on molecular data in terms of providing the basic

phylogenetic framework for angiosperms morphological and molecular data are only

very rarely in irreconcilably strong conflict There are indeed a few places where the

conflict seems extreme These include the relative positions of the Monimiaceae and

Hernandiaceae (Laurales) the position of Hanguanaceae (Commelinales [as here] or

Zingiberales) and of Triplostegia (is it in Dipsacaceae or Valerianaceae - see

Dipsacales) Fossils are unlikely to affect the topologies of the trees presented here but

see below for their importance in understanding morphological evolution in general and

the evolution of angiosperms in particular Nevertheless some largely reject the idea

that trees based on molecular data alone can recover phylogenetic relationships

especially when branching points are old and prefer to used trees based on analysis of

morphological data including those taken from fossils (Hilton amp Bateman 2006 Farjon

2007)

All in all however the extent of the congruence between morphological and

molecular data is impressive and heartening and many clades can be characterised

morphologically It seemed in 1998 that there were no unambiguous morphological

synapomorphies for angiosperm orders (K Bremer 2000) and this is still true if by

unambiguous is meant non-homoplasious However many orders and other clades

have synapomorphies even if these may be indistinguishable at least at the current

level of morphological and developmental knowledge from parallel occurences

elsewhere As our morphological knowledge increases so too does the number of

apomorphies

Endress and Matthews (2006a) emphasize the importance of tendencies and

developmental constraints when thinking about characters of clades within the rosids -

many characters or character combinations occur in a rather sporadic fashion within

the clade and apparently notably less frequently outside it There are a number of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

28

examples of tendencies here and in the individual order pages such as the features

enclosed by parentheses in groups above the level of family eg see (cuticular waxes

as aggregated rodlets) for the commelinids and the discussion of the distribution of

polyandry that is flowers with numerous stamens in the asterid I + II groups etc

Tendencies may involve more than single characters When thinking of the

associations of features like integument thickness and vascularization of the integument

(Endress amp Matthews 2006a) one can imagine fairly simple developmental

preconditions being operative As Endress and Matthews (2006a) note it is difficult to

imagine vascular tissue developing in an integument only two cells thick Changes here

would be loosely correlated if morphologically linked However the strongly correlated

changes noted by Givnish et al (2005) are ecologically linked but are presumably

morphologicallydevelopmentally independent When there seem to be characters

evolving more or less together tests can be carried out to see if the changes are

concentrated on certain branches of the tree (eg Maddison 1990 Sanderson 1991

Maddison amp Maddison 2000) Such tests have rarely been carried out at higher levels in

angiosperms

One of the most striking examples of a tendency is the distribution of N-fixation

restricted as it is to a monophyletic group of four clades although it has arisen seven (or

perhaps several more) times independently within the N-fixing clade and several

members of two quite different kinds of bacteria are involved (eg Clawson et al 2004

Elliott et al 2007 Sprent amp James 2007) Other examples of work that bears on the issue

of tendencies include the findings that flowers of polysymmetrical Arabidopsis have

genes like TCP1 that are expressed asymmetrically during early development and TCP1

is a probable orthologue of the well-known CYC gene of Antirrhinum that is involved in

the development of monosymmetric slowers there (Cubas et al 2001 Costa et al 2005

etc) Parallelism might build on this underlying morphologically cryptic monosymmetry

even if details of the genetic mechanisms causing the monosymmetry evident in

particular groups may be different (Reeves amp Olmstead 2003 Cubas 2004 see above)

Similarly there have been several recent suggestions that the capability to synthesise a

particular metabolite may be switched off but not lost and so can sometimes be

reacquired (eg Wink amp Witte 1983 Wink 2003 Liscombe et al 2005 Larsson 2007

Waterman 2007) Hence perhaps the rather spotty distribution of many secondary

metabolites like ellagic acid the indole alkaloid camptothecin iridoids etc when

considered in the context of phylogenies As a non-botanical example - but a rather nice

one - Salwini-Plawen and Mayr (1961) suggested some time ago that there has been

considerable parallelism (40-65 or more independent origins) in the evolution of eyes in

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

29

metazoans However Pax 6 seems to be a master control gene that is involved in eye

formation perhaps throughout all bilateralians (eg Gehring amp Ikeo 1999 Erwin amp

Davidson 2002) which changes the nature of the problem Note however that in

general our invocation of evolutionary tendencies to explain the patterns of

distributions of characters is really an admission that we do not understand those

patterns

As we find out more about variation we will probably find fewer and fewer features

constant throughout a group Most unqualified statements of presence and absence

should properly be qualified as usually present or usually absent if one is thinking of

the characterisations on the order pages as encompassing the total variation within a

clade Thus Pistia alone among monocots as so far known has sieve tube plastids with

starch grains not protein crystals However this does not affect the fact that sieve tube

plastids with cuneate protein crystals and lacking starch grains are an apomorphy for

monocots Furthermore fossils in a number of cases suggest character combinations

unknown in extant taxa as may be seen in the discussions of Fagaceae Platanaceae

Iteaceae Calycanthaceae etc Confusing the issue there may be questions as to where

exactly on the tree a particular fossil is to be placed (eg see Nymphaeaceae

Calycanthaceae Archaefructus etc)

To summarize given our current understandings of both phylogenies and

characters evolution of some characters in which we are interested seems very labile

(see eg D Soltis et al 2005b Endress and Matthews 2006a Stevens 2006b Ekman et

al 2008 for a good recent study) and I have been cautious when talking about character

evolution Much effort must continue to be spent in summarizing characters of well-

established clades at all levels providing features by which they may be recognized and

signaling synapomorphies Remember that (1) the basic morphological anatomical and

chemical knowledge of many critical taxa is woefully incomplete (2) different

assumptions about character evolution may greatly affect the position of

synapomorphies on trees (3) in many cases relationships within and between many

groups are too uncertain at present to worry very much about synapomorphies and

(4) we must be clear about what we do and do not not know As mentioned above I

have indicated in a number of places where there are particularly important

uncertainties as to where characters should be placed on the tree Nevertheless it is a

relatively easy matter to update notes such as these and it can be a simple matter to

incorporate new data on characters that have never before been considered in the

context of a tree Much basic - and unfortunately perhaps unfashionable - work must be

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

30

carried to clarify the distribution of morphological anatomical and chemical characters

for examples of what can to be done see the work of P K Endress and collaborators

(floral morphology and development) and S R Jensen and collaborators (iridoids) But

acquiring information about nodal anatomy cell and tissue distribution nodal anatomy

and the like is not difficult at all and should be generally encouraged

But all this is merely a necesary prelude to the understanding of evolution For this

one needs to know a lot more including dating the phylogeny understanding the

function(s) of characters factoring in relevant aspects of the palaeoenvironment in

which an apomorphy first appeared etc

SUMMARY OF THE SYSTEM

Below is a formalised summary of the relationships within orders of the families of seed

plants There are a few families that are not recognised even as options in APG II and

vice versa as well as a few extra orders The families to be recognised in the most

recent edition of Mabberleys The Plant Book (Mabberley 2008) are also largely

consistent with those below But all differences are trivial and will - I hope - eventually

disappear

Square brackets - [] - enclose clades the plus sign - + - designates sister taxa a

comma - - denotes part of a polytomy and quotation marks - - denotes a

paraphyletic group

SEED PLANTS

GYMNOSPERMS

Cycadales

Cycadaceae + Zamiaceae

Ginkgoales

Ginkgoaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

31

Pinales

Pinaceae [[Araucariaceae [Podocarpaceae + Phyllocladaceae]] [Sciadopityaceae

[Taxaceae + Cupressaceae]]]

Gnetales

Ephedraceae [Gnetaceae + Welwitschiaceae]

ANGIOSPERMSFLOWERING PLANTS

Amborellales

Amborellaceae

Nymphaeales

Hydatellaceae [Cabombaceae + Nymphaeaceae]

Austrobaileyales

Austrobaileyaceae [Schisandraceae + Trimeniaceae]

Chloranthales

Chloranthaceae

MAGNOLIIDS

Magnoliales

Myristicaceae [Magnoliaceae [[Himantandraceae + Degeneriaceae] [Eupomatiaceae +

Annonaceae]]]

Laurales

Calycanthaceae [[Siparunaceae [Gomortegaceae + Atherospermataceae]] [Monimiaceae

[Hernandiaceae + Lauraceae]]]

Canellales

Canellaceae + Winteraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

32

Piperales

[Hydnoraceae + Aristolochiaceae] [Piperaceae + Saururaceae]

MONOCOTS

Acorales

Acoraceae

Alismatales

Araceae [Tofieldiaceae [[Alismataceae [Hydrocharitaceae + Butomaceae]]

[Scheuchzeriaceae [Aponogetonaceae [Juncaginaceae [Maundiaceae [[Posidoniaceae

[Ruppiaceae + Cymodoceaceae]] [Zosteraceae + Potamogetonaceae]]]]]]]

Petrosaviales

Petrosaviaceae

Dioscoreales

Nartheciaceae [[Taccaceae + Thismiaceae] [Burmanniaceae + Dioscoreaceae]]

Pandanales

Velloziaceae Triuridaceae Stemonaceae [Pandanaceae + Cyclanthaceae]

Liliales

Corsiaceae [Campynemataceae [Petermanniaceae [Colchicaceae + Alstroemeriaceae]]

Melanthiaceae [[Philesiaceae + Rhipogonaceae] [Smilacaceae + Liliaceae]]]

Asparagales

Orchidaceae [[Boryaceae [Blandfordiaceae [Lanariaceae [Asteliaceae + Hypoxidaceae]]]]

[[Ixioliriaceae + Tecophilaeaceae] [Doryanthaceae [Iridaceae [Xeronemataceae

[Xanthorrhoeaceae [Alliaceae + Asparagaceae]]]]]]]

COMMELINIDS

Unplaced

Dasypogonaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

33

Arecales

Arecaceae

Poales

[Typhaceae + Bromeliaceae] [Rapateaceae [[Thurniaceae [Juncaceae + Cyperaceae]]

[[Mayacaceae [Eriocaulaceae + Xyridaceae]] [Flagellariaceae [Anarthriaceae

[Centrolepidaceae + Restionaceae]]] [Joinvilleaceae [Ecdeiocoleaceae + Poaceae]]]]]]]

Commelinales

[Commelinaceae + Hanguanaceae] [Philydraceae [Haemodoraceae + Pontederiaceae]]

Zingiberales

Musaceae [Strelitziaceae + Lowiaceae] Heliconiaceae [[Cannaceae + Marantaceae]

[Costaceae + Zingiberaceae]]

Ceratophyllales

Ceratophyllaceae

EUDICOTS

Ranunculales

Eupteleaceae [Papaveraceae [[[Lardizabalaceae + Circaeasteraceae] [Menispermaceae

[Berberidaceae + Ranunculaceae]]]]

Sabiales

Sabiaceae

Proteales

Nelumbonaceae [Platanaceae + Proteaceae]

Trochodendrales

Trochodendraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

34

Buxales

Haptanthaceae [Buxaceae + Didymelaceae]

CORE EUDICOTS

Gunnerales

Gunneraceae + Myrothamnaceae

Dilleniales

Dilleniaceae

Saxifragales

Peridiscaceae [[Paeoniaceae [Altingiaceae [Hamamelidaceae [Cercidiphyllaceae +

Daphniphyllaceae]]]] [[Crassulaceae [Aphanopetalaceae [Tetracarpaeaceae

[Penthoraceae + Haloragaceae]]]] [Iteaceae [Grossulariaceae + Saxifragaceae]]]]

Cynomoriaceae unplaced

Vitales

Vitaceae

ROSIDS

FABIDROSID I

Zygophyllales

Krameriaceae + Zygophyllaceae

Celastrales

Lepidobotryaceae + Celastraceae

Oxalidales

Huaceae [[Connaraceae + Oxalidaceae] [Cunoniaceae [Elaeocarpaceae [Brunelliaceae +

Cephalotaceae]]]]

Malpighiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

35

[Achariaceae [Goupiaceae [Violaceae + Passifloraceae] [Lacistemataceae Salicaceae]]

[Lophopyxidaceae Putranjivaceae] [Ctenolophonaceae [Erythroxylaceae +

Rhizophoraceae]] Linaceae Ixonanthaceae Humiriaceae Irvingiaceae

Centroplacaceae Pandaceae [[Bonnetiaceae + Clusiaceae] [Calophyllaceae

[Hypericaceae + Podostemaceae]]] [Malpighiaceae + Elatinaceae] Ochnaceae

[Peraceae [Rafflesiaceae + Euphorbiaceae]] [Phyllanthaceae + Picrodendraceae]

[Balanopaceae [[Trigoniaceae + Dichapetalaceae] [Chrysobalanaceae +

Euphroniaceae]]] Caryocaraceae

N-FIXING CLADE

Fabales

Quillajaceae [Fabaceae [Polygalaceae + Surianaceae]]

Rosales

Rosaceae [[Barbeyaceae [Elaeagnaceae Dirachmaceae Rhamnaceae]] [Ulmaceae

[Cannabaceae [Moraceae + Urticaceae]]]]

Cucurbitales

Anisophylleaceae [[Corynocarpaceae + Coriariaceae] [Cucurbitaceae [Tetramelaceae

[Datiscaceae + Begoniaceae]]]] Apodanthaceae

Fagales

Nothofagaceae [Fagaceae [[Myricaceae + Juglandaceae] [Casuarinaceae

[Ticodendraceae + Betulaceae]]]]

MALVIDROSID II

pgtGeraniales

Geraniaceae [[Melianthaceae + Francoaceae] [Vivianaceae + Ledocarpaceae]]

Myrtales

Combretaceae [[Onagraceae + Lythraceae] [[Vochysiaceae + Myrtaceae]

[Melastomataceae [Crypteroniaceae [Alzateaceae + Penaeaceae]]]]]

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

36

Crossosomatales

[Staphyleaceae [Guamatelaceae [Crossosomataceae + Stachyuraceae]]] [Aphloiaceae

[Geissolomataceae + Strasburgeriaceae]]

Picramniales

Picramniaceae

Sapindales

Biebersteiniaceae Nitrariaceae [Kirkiaceae [Anacardiaceae + Burseraceae]]

[Sapindaceae [Simaroubaceae Rutaceae Meliaceae]]

Huerteales

Gerradinaceae [Dipentodontaceae + Tapisciaceae]

Malvales

Neuradaceae [Thymelaeaceae [Sphaerosepalaceae Bixaceae [Cistaceae

[Sarcolaenaceae + Dipterocarpaceae]] [Cytinaceae + Muntingiaceae] Malvaceae]]

Brassicales

[Akaniaceae + Tropaeolaceae] [[Moringaceae + Caricaceae] [Setchellanthaceae

[Limnanthaceae [[Koeberliniaceae [Bataceae + Salvadoraceae]] [Emblingiaceae

[Pentadiplandraceae [Gyrostemonaceae Resedaceae Stixis and relatives] Tovariaceae

[Capparaceae [Cleomaceae + Brassicaceae]]]]]]]]

Berberidopsidales

Aextoxicaceae + Berberidopsidaceae

Santalales

Erythropalaceae Olacaceae [[Loranthaceae [Misodendraceae + Schoepfiaceae]]

[Opiliaceae + Santalaceae]] Balanophoraceae unplaced

Caryophyllales

[[Droseraceae [Nepenthaceae [Drosophyllaceae [Ancistrocladaceae +

Dioncophylleaceae]]]]] [[Frankeniaceae + Tamaricaceae] [Polygonaceae +

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

37

Plumbaginaceae]]] [Rhabdodendraceae [Simmondsiaceae [[Asteropeiaceae +

Physenaceae] [[Caryophyllaceae [Achatocarpaceae + Amaranthaceae]]

[Stegnospermataceae [Limeaceae [[Lophocarpaceae [Barbeuiaceae [Aizoaceae

[Sarcobataceae Phytolaccaceae Nyctaginaceae]]]] [Molluginaceae [Halophytaceae

Montiaceae Basellaceae Didiereaceae [Talinaceae [Portulacaceae [Anacampseros etc +

Cactaceae]]]]]]]]]]]]]

ASTERIDS

Cornales

[Cornaceae + Nyssaceae] [Hydrangeaceae + Loasaceae] Hydrostachyaceae [Curtisiaceae

+ Grubbiaceae]

Ericales

[Balsaminaceae [Marcgraviaceae + Tetrameristaceae]] [[Polemoniaceae +

Fouquieraceae] Lecythidaceae [[Sladeniaceae + Pentaphylacaceae] [Sapotaceae

[Ebenaceae [Maesaceae [Theophrastaceae [Myrsinaceae + Primulaceae]]]]]

[Mitrastemonaceae Theaceae [Symplocaceae [Styracaceae + Diapensiaceae]]

[[Sarraceniaceae [Roridulaceae + Actinidiaceae] [Clethraceae [Cyrillaceae +

Ericaceae]]]]]]

LAMIIDASTERID I

Unplaced

Metteniusaceae Oncothecaceae Icacinaceae and some unplaced ex-Icacinaceae

Garryales

Garryaceae + Eucommiaceae

Unplaced

Boraginaceae Vahliaceae

Gentianales

Rubiaceae [Gentianaceae [Loganiaceae [Gelsemiaceae + Apocynaceae]]]

Lamiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

38

Plocospermataceae [Oleaceae [Tetrachondraceae [Calceolariaceae

[Gesneriaceae Peltanthera [Pedaliaceae Carlemanniaceae Martyniaceae Stilbaceae

Plantaginaceae Linderniaceae Bignoniaceae Verbenaceae Lamiaceae Paulowniaceae

Schlegeliaceae Thomandersiaceae Phrymaceae Rehmannia Acanthaceae

Scrophulariaceae Orobanchaceae Byblidaceae Lentibulariaceae]]]]]

Solanales

[Montiniaceae [Sphenocleaceae + Hydroleaceae]] [Convolvulaceae + Solanaceae]

CAMPANULIDASTERID II

Aquifoliales

[[Cardiopteridaceae + Stemonuraceae] [Aquifoliaceae [Helwingiaceae +

Phyllonomaceae]]

Asterales

[Rousseaceae + Campanulaceae] Pentaphragmataceae [[Alseuosmiaceae [Phellinaceae

+ Argophyllaceae]] Stylidiaceae [Menyanthaceae [Goodeniaceae [Calyceraceae +

Asteraceae]]]]

Escalloniales

Escalloniaceae

Bruniales

[Bruniaceae + Columelliaceae]

Apiales

Pennantiaceae [Torricelliaceae [Griseliniaceae [Pittosporaceae [Araliaceae

[Myodocarpaceae + Apiaceae]]]]]

Paracryphiales

Paracryphiaceae

Dipsacales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

39

Adoxaceae [Diervillaceae [Caprifoliaceae [Linnaeaceae [Morinaceae [Dipsacaceae +

Valerianaceae]]]]]

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MAIN GROUPINGS

Asterids commelinids core eudicots asterid 1 asterid

2 eudicots gymnosperms Magnoliophyta monocots N-fixing claderosids seed plants

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Abietales Acanthales Acerales Acorales Actinidiales Actinostrobales Adoxales Aescul

ales Agavales Aizoales AkanialesAlismatales Alliales Alseuosmiales Alstroemeriales

Altingiales Amaranthales Amaryllidales Amborellales AmbrosialesAmmiales Amomal

es Ancistrocladales Anisophylleales Annonales Anthobolales Apiales Apocynales Apo

nogetonalesAquifoliales Arales Araliales Aralidiales Araucariales Arecales Aristolochi

ales Athrotaxidales Asarales AsclepiadalesAsparagales Asphodelales Asparagales Ast

eliales Atriplicales Aucubales Austrobaileyales Avenales

Balanitales Balanopales Balanophorales Balsaminales Barbeyales Barclayales Batales

Begoniales BerberidalesBerberidopsidales Betulales Biebersteiniales Bignoniales Bix

ales Boraginales Brassicales Brexiales Bromeliales BrunialesBrunoniales Burmannial

es Burserales Butomales Buxales Byblidales

Cactales Callitrichales Calycanthales Calycerales Campanulales Campynematales Can

ellales Cannales CapparalesCaprifoliales Cardiopteridales Carduales Caricales Carle

manniales Caryophyllales Cassiales Casuarinales CelastralesCentrolepidales Cephalot

ales Cephalotaxales Ceratophyllales Cercidiphyllales Chenopodiales Chironiales Chlor

anthalesChrysobalanales Cinchonales Circaeasterales Cistales Citrales Cocosales Col

chicales Columelliales CombretalesCommelinales Connarales Convolvulales Coriarial

es Cornales Corylales Corynocarpales Crassulales CrossosomatalesCucurbitales Cunn

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

40

inghamiales Cunoniales Cupressales Cycadales Cyclanthales Cymodoceales Cynarales

CynomorialesCyperales Cyrillales Cytinales

Daphnales Daphniphyllales Dasypogonales Datiscales Degeneriales Desfontainiales D

iapensiales Didymelales DillenialesDioales Dioncophyllales Dioscoreales Diospyrales

Dipentodontales Dipsacales Droserales

Ebenales Echiales Elaeagnales Elaeocarpales Elatinales Elodeales Emmotales Empetr

ales Ephedrales Ericales EriocaulalesErythropalaless Escalloniales Eucommiales Euph

orbiales Eupomatiales Eupteleales Euryalaless

Fabales Fagales Falcatifoliales Ficales Flacourtiales Flagellariales Fouquieriales Franc

oales Frangulales

Galiales Garryales Geissolomatales Gentianales Geraniales Gesneriales Ginkgoales G

laucidiales Globulariales GnetalesGoodeniales Greyiales Griseliniales Grossulariales

Grubbiales Gunnerales Gyrocarpales Gyrostemonales

Haemodorales Haloragales Hamamelidales Hanguanales Heisteriales Helleborales He

lwingiales Himantandrales HippuridalesHomaliales Hortensiales Huales Huerteales

Hydatellales Hydnorales Hydrangeales Hydrastidales HydrocharitalesHydropeltidales

Hydrostachyales Hypericales Hypoxidales

Icacinales Illiciales Iridales Irvingales Iteales Ixiales

Jasminales Juglandales Julianiales Juncaginales Juncales

Lacistematales Lactoridales Lamiales Lardizabalales Laurales Lecythidales Ledocarpal

es Leitneriales LentibularialesLigustrales Liliales Limnanthales Linales Liriales Loasal

es Lobeliales Loganiales Lonicerales Loranthales LowialesLythrales

Magnoliales Malpighiales Malvales Marathrales Marcgraviales Mayacales Medusagy

nales Medusandrales MelanthialesMelastomatales Meliales Melianthales Meliosmal

es Menispermales Menyanthales Metteniusales MitrastemonalesMiyoshiales Moni

miales Moringales Musales Myricales Myristicales Myrothamnales Myrsinales Myrta

les

Najadales Nandinales Narcissales Nartheciales Nelumbonales Nepenthales Neuradal

es Nitrariales Nolanales NothofagalesNyctaginales Nymphaeales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Ochnales Oenotherales Olacales Oleales Onagrales Oncothecales Opuntiales Orchid

ales Oxalidales

Saxifragales Pandales Pandanales Papaverales Paracryphiales Parasitaxales Paridales

Parnassiales PassifloralesPenaeales Pennantiales Pentaphragmatales Peridiscales P

etiveriales Petrosaviales Phellinales Philydrales PhyllanthalesPhysenales Phytolaccal

es Picramniales Pinales Pinguiculales Piperales Pittosporales Plantaginales Platanale

sPlumbaginales Poales Podocarpales Podophyllales Podostemales Polemoniales Pol

ygalales Polygonales PontederialesPortulacales Posidoniales Potamogetonales Primu

lales Proteales

Quercales Quillajales Quintiniales

Rafflesiales Ranunculales Rapateales Resedales Restionales Rhabdodendrales Rhamn

ales Rhinanthales RhizophoralesRhodorales Rhoipteleales Roridulales Rosales Rouss

eales Rubiales Ruppiales Rutales

Sabiales Salicales Salvadorales Samolales Samydales Sanguisorbales Santalales Sapin

dales Sapotales SarracenialesSaxegotheales Saxifragales Scheuchzeriales Sciadopitya

les Scleranthales Scrophulariales Scyphostegiales SedalesSilenales Simmondsiales S

milacales Solanales Sphenocleales Sphenostemonales Stangeriales Stellariales Stemo

nalesStilbales Stylidiales Styracales Surianales

Taccales Tamales Tamaricales Taxales Taxodiales Tecophilaeales Terebinthales Tern

stroemiales Theales TheligonalesThymelaeales Tiliales Tofieldiales Torricelliales Tov

ariales Tribelales Trilliales Trimeniales Triuridales TrochodendralesTropaeolales Turn

erales Typhales

Ulmales Urticales

Vacciniales Vahliales Vallisneriales Velloziales Veratrales Verbenales Viburnales Vinc

ales Violales Viscales VitalesVochysiales

Welwitschiales Winterales

Xanthorrhoeales Ximeniales Xyridales

Zamiales Zingiberales Zosterales Zygophyllales

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which the genus of the basionym is to be found

Abietaceae Abolbodaceae Abrophyllaceae Acalyphaceae Acanthaceae [Acanthochla

mydaceae] Acarnaceaee AceraceaeAchariaceae Achatocarpaceae Achradaceae Achy

ranthaceae Acmopylaceae Acoraceae Acristaceae ActaeaceaeActinidiaceae Actinota

ceae Amentotaxaceae Adenogrammaceae Adoxaceae Aegialitidaceae Aegicerataceae

Aegilopaceae Aeginetiaceaee Aegiphilaceae Aesculaceae Aextoxicaceae Agapanthac

eae Agavaceae AgdestidaceaeAgialidaceae Agrimoniaceae Agrostidaceae Ailanthace

ae Aitoniaceae Aizoaceae Akaniaceae Alangiaceae AlchemillaceaeAldrovandaceae A

lismataceae Alliaceae Allioniaceae Allophylaceae Aloaceae Alopecuraceae Alpiniacea

e AlseuosmiaceaeAlsinaceae Alsodeiaceae Alstroemeriaceae Altingiaceae Alzateacea

e Amaranthaceae Amaryllidaceae Amborellaceae Ambrosiaceae Amentotaxaceae A

mmanniaceae Ammiaceae Amomaceae Ampelidopsaceae Amygdalaceae Amyridacea

eAnacardiaceae Anagallidaceae Anarthriaceae Anchusaceae Ancistrocladaceae Andr

omedaceae AndropogonaceaeAndrostachyaceae Androsynaceae Anemarrhenaceae

Anemonaceae Angelicaceae Anisophylleaceae AnnonaceaeAnomochloaceae Anopter

aceae Anrederaceae Anthemidaceae Anthericaceae Anthobolaceae Antidesmataceae

Antirrhinaceae Antoniaceae Aparinaceae Aphanopetalaceae Aphloiaceae Aphyllanth

aceae Apiaceae ApocynaceaeApodanthaceae Aponogetonaceae Aporusaceae Apose

ridaceae Apostasiaceae Aptandraceae AquifoliaceaeAquilariaceae Aquilegiaceae A

raceae Aragoaceae Araliaceae Aralidiaceaee Araucariaceae Arbutaceae Arceuthidace

aeArceuthobiaceae Arctostaphylaceae Arctotidaceae Ardisiaceae Arecaceae Argoph

yllaceae Arisaraceae AristolochiaceaeAristoteliaceae Arjonaceae Armeriaceae Artem

isiaceae Arthrotaxidaceae Artocarpaceae Arundinaceae ArundinellaceaeAsaraceae A

sclepiadaceae Ascyraceaee Asiraceae Asparagaceae Asperulaceae Asphodelaceae As

pidistraceaeeAsteliaceae Asteraceae Asteranthaceae Asteropeiaceae Astragalaceae

Astrocarpaceae AthanasiaceaeAtherospermataceae Atriplicaceae Atropaceae Aucuba

ceae Aurantiaceae Austrobaileyaceae Austrotaxaceae AvenaceaeAverrhoaceae Avet

raceae Avicenniaceae Azaleaceae Azimaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Balanitaceae Balanopaceae Balanophoraceae Balsameaceae Balsaminaceae Bambusa

ceae Barbaceniaceae BarbeuiaceaeBarbeyaceae Barclayaceae Barringtoniaceae Base

llaceae Bataceae Baueraceae Bauhiniaceae Baxteriaceae BegoniaceaeBehniaceae B

elangeraceae Belloniaceae Bembiciaceae Berberidaceae Berberidopsidaceae Berryac

eaee BersamaceaeBertyaceae Berzeliaceae Besleriaceae Betaceae Betulaceae Biebe

rsteiniaceae Bifariaceae Bignoniaceae BischofiaceaeBixaceae Blakeaceae Blandfordia

ceae Blattiaceae Blepharocaryaceae Blitaceae Blyxaceae Boerlagellaceae Bolivaracea

eBombacaceae Bonnetiaceae Bontiaceae Boopidaceae Boraginaceae Borassaceae B

oroniaceae BoryaceaeBotryodendraceae Bougainvilleaceae Boweniaceae Brassicacea

e Brachycaulaceae Bretschneideraceae BrexiaceaeBromeliaceae Brownlowiaceae Br

unelliaceae Bruniaceae Brunoniaceae Brunsvigiaceae Bryoniaceae BuchneraceaeBuci

daceae Buddlejaceae Buglossaceae Bulbocodiaceae Bumeliaceae Bupleuraceae Burc

hardiaceae BurmanniaceaeBurseraceae Butneriaceae Butomaceae Buxaceae Byblida

ceae Byttneriaceae

Cabombaceae Cacaoaceae Cactaceae Caesalpiniaceae Caladiaceae Calamaceae Calce

olariaceae CalectasiaceaeCalendulaceae Callaceae Callicomaceae Calligonaceae Calli

traceae Callitrichaceae Calochortaceae CalophyllaceaeCalthaceae Calycanthaceae Ca

lyceraceae Cambogiaceae Camelliaceae Campanulaceae CampynemataceaeCanacom

yricaceae Canellaceae Cannabaceae Cannaceae Canopodaceae Canotiaceae Cansjera

ceae CapparaceaeCaprariaceae Caprifoliaceae Cardiopteridaceae Carduaceae Carica

ceae Carissaceae Carlemanniaceae CarpinaceaeCarpodetaceae Cartonemataceae Ca

ryocaraceae Caryophyllaceae Casearieaceae Cassiaceae CassipoureaceaeCassythacea

e Castaneaceae Castelaceae Casuarinaceae Catesbaeaceae Cathedraceae Cecropiace

ae CedraceaeCedrelaceae Celastraceae Celosiaceae Celtidaceae Centaureaceae Cen

trolepidaceae Centroplacaceae CepaceaeCephalanthaceae Cephalotaceae Cephalota

xaceae Cerastiaceae Ceratoniaceae Ceratophyllaceae CerberaceaeCercidiphyllaceae

Cercocarpaceae Cercodiaceae Cereaceae Cerinthaceae Ceroxylaceae Cestraceae Cev

alliaceaeChailletiaceae Chamaedoraceae Chamaemoraceae Chamelauciaceae Chaun

ochitonaceae Chelidoniaceae ChelonaceaeChenopodiaceae Chimonanthaceae Chingi

thamnaceae Chionographidaceae Chiranthodendraceae ChironiaceaeChlaenaceae Ch

loanthaceae Chloranthaceae Chloridaceae Chlorogalaceae Chrysobalanaceae Cicerac

eae CichoriaceaeCimicifugaceae Cinchoniaceae Circaeaceae Circaeasteraceae Cissac

eae Cistaceae Citraceae Clematidaceae CleomaceaeClethraceae Clusiaceae Cneorac

eae Cnestidaceae Cnicaceae Cobaeaceae Cochlospermaceae Cocosaceae Coffeaceae

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Colchicaceae Coleogynaceae Columelliaceae Combretaceae Commelinaceae Comocl

adiaceae Compositae CompsoaceaeConantheraceae Connaraceae Conostylidaceae C

onvallariaceae Convolvulaceae Cordiaceae CoreopsidaceaeCoriandraceae Coriariacea

e Coridaceae Corispermaceae Cornaceae Corokiaceae Coronillaceae Correaceae Cor

rigiolaceaeCorsiaceae Corydalaceae Corylaceae Corynocarpaceae Coryphaceae Cost

aceae Cotyledonaceae CoulaceaeCoutariaceae Coutoubiaceae Crassulaceae Crescen

tiaceae Cressaceae Crinaceae Crocaceae CroomiaceaeCrossosomataceae Crotonacea

e Cruciferae Cryptaceae Crypteroniaceae Cryptocorynaceae CryptomeriaceaeCtenol

ophonaceae Cucurbitaceae Cunninghamiaceae Cunoniaceae Cupressaceae Curcumac

eae Curtisiaceae CuscutaceaeCyananthaceae Cyanastraceae Cynanchaceae Cyanella

ceae Cycadaceae Cyclanthaceae CyclantheraceaeCyclocheilaceae Cydoniaceae Cymo

doceaceae Cynaraceae Cynocrambaceae Cynomoriaceae Cyperaceae CyphiaceaeCyp

hocarpaceae Cypripediaceae Cyrillaceae Cyrtandraceae Cyrtanthaceae Cytinaceae

Dacrycarpaceae Dactylanthaceae Damasoniaceae Daphnaceae Daphniphyllaceae Das

ypogonaceae DatiscaceaeDaturaceae Daucaceae Davidiaceae Davidsoniaceae Decais

neaceae Deeringiaceae Degeneriaceae DelphiniaceaeDendrophthoaceae Columelliac

eae Detariaceae Dialypetalanthaceae Dianellaceae Dianthaceae DiapensiaceaeDicha

petalaceae Dichondraceae Diclidantheraceae Dicrastylidiaceae Dictamnaceae Didiere

aceae DidymelaceaeDidymocarpaceae Diegodendraceae Diervillaceae Digitalidaceae

e Dilatridaceae Dilleniaceae Dioaceae DionaeaceaeDioncophyllaceae Dioscoreaceae

Diosmaceae Diospyraceae Dipentodontaceae Diphylleiaceae Diplolaenaceae Dipsaca

ceaeDipterocarpaceae Dirachmaceae Disanthaceae Diselmaceae Dodonaeaceae Do

mbeyaceae Donatiaceae DorsteniaceaeDortmannaceae Doryanthaceae Drabaceae D

racaenaceae Dracontiaceae Drimyidaceae Droseraceae DrosophyllaceaeDryadaceae

Duabangaceae Duckeodendraceae Dulongiaceae Durantaceae Durionaceae Dysphani

aceae

Ebenaceae Eccremidaceae Ecdeiocoleaceae Echiaceae Echinopaceae Ehretiaceae Ela

eagnaceae ElaeocarpaceaeElegiaceae Elatinaceae Ellisiaceae Ellisiophyllaceae Elodea

ceae Elytranthaceae Embeliaceae Emblingiaceae EmmotaceaeEmpetraceae Encepha

lartaceae Engelhardtiaceae Enhalaceae Epacridaceae Ephedraceae Ephemeranthacea

eEpilobiaceae Epimediaceae Eragrostidaceae Eremolepidaceae Eremosynaceae Erica

ceae Erinaceae EriocaulaceaeEriogonaceae Eriospermaceae Erodiaceae Erycibaceae

Eryngiaceae Erysimaceae Erythroniaceae ErythropalaceaeErythrospermaceae Erythro

xylaceae Escalloniaceae Eschscholziaceae Eucomidaceae Eucommiaceae Eucryphiace

aeEuonymaceae Eupatoriaceae Euphorbiaceae Euphrasiaceae Euphroniaceae Eupo

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matiaceae Eupteleaceae EuryalaceaeEustrephaceae Euthemidaceae Eutocaceae Exb

ucklandiaceae Exocarpaceae

Fabaceae Fagaceae Falcatifoliaceae Ferulaceae Festucaceae Ficaceae Ficoidaceae Fit

zroyaceae FlacourtiaceaeFlagellariaceae Flindersiaceae Foetidiaceae Forestieraceae

Fothergillaceae Fouquieriaceae Fragariaceae FrancoaceaeFrangulaceae Frankeniacea

e Fraxinaceae Fraxinellaceae Freycinetiaceae Fritillariaceae Fuchsiaceae Fumariacea

eFunkiaceae

Gaiadendraceae Galacaceae Galanthaceae Galaxiaceae Galedupaceae Galeniaceae G

aliaceae Garciniaceae GardeniaceaeGardneriaceae Garryaceae Geissolomataceae Ge

itonoplesiaceae Gelsemiaceae Geniostomaceae GentianaceaeGeonomataceae Geosir

idaceae Geraniaceae Gerrardinaceae Gesneriaceae Gethyllidaceae Gilliesiaceae Gina

lloaceaeGinkgoaceae Gisekiaceae Gladiolaceae Glaucidiaceae Glechomaceae Glinace

ae Globulariaceae Gnaphaliaceae GnetaceaeGoetzeaceae Gomortegaceae Gomphiac

eae Gomphrenaceae Gonystylaceae Goodeniaceae Gordoniaceae GouaniaceaeGoup

iaceae Gramineae Gratiolaceae Grewiaceae Greyiaceae Grielaceae Griseliniaceae Gr

onoviaceae GrossulariaceaeGrubbiaceae Guaiacanaceae Guamatelaceae Guettardac

eae Gunneraceae Gustaviaceae Guttiferae GyrocarpaceaeGyrostemonaceae

Hachetteaceae Haemanthaceae Haemodoraceae Halesiaceae Hallieraceae Halocarpa

ceae Halophilaceae HalophytaceaeHaloragaceae Hamamelidaceae Hanguanaceae H

aptanthaceae Harmandaceae Hebenstretiaceae HectorellaceaeHederaceae Hedyosm

aceae Hedyotidaceae Hedysaraceae Heisteriaceae Heleniaceae Heliamphoraceae H

elianthaceaeHelianthemaceae Helichrysaceae Heliconiaceae Helicteraceae Heliotropi

aceae Helleboraceae Heloniadaceae HelosidaceaeHelwingiaceae Hemerocallidaceae

Hemimeridaceae Hemiphylacaceae Henriqueziaceae Henslowiaceae Hermanniaceae

Hernandiaceae Herniariaceae Herreriaceae Hesperocallidaceae Heterantheraceae He

teropyxidaceae HeterostylaceaeHibbertiaceae Hibiscaceae Hilleriaceae Himantandra

ceae Hippocastanaceae Hippocrateaceae HippomanaceaeHippophaeaceae Hippurida

ceae Hirtellaceae Holacanthaceae Homaliaceae Hopkinsiaceae Hoplestigmataceae H

ordeaceaeHornschuchiaceae Hortensiaceae Hortoniaceae Hostaceae Houstoniaceae

Huaceae Huerteaceae HugoniaceaeHumbertiaceae Humiriaceae Hyacinthaceae Hyd

atellaceae Hydnoraceae Hydrangeaceae Hydrastidaceae HydrillaceaeHydrocharitacea

e Hydrocotylaceae Hydrogetonaceae Hydroleaceae Hydropeltidaceae Hydrophylacac

eae HydrophyllaceaeHydrostachyaceae Hymenocardiaceae Hypecoaceae Hyperanth

eraceae Hypericaceae Hypopityaceae HypoxidaceaeHypseocharitaceae

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Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta

ceae ImperatoriaceaeInocarpaceae Inulaceae Ionidiaceae Iriarteaceae Iridaceae Irvi

ngiaceae Isnardiaceae Isophysidaceae IteaceaeStrasburgeriaceae Ixiaceae Ixioliriace

ae Ixonanthaceae

Jalapaceae Jamboliferaceae Japonoliriaceae Jasionaceae Jasminiaceae Johnsoniaceae

Joinvilleaceae JuglandaceaeJulianiaceae Juncaceae Juncaginaceae Juniperaceae Jus

siaeaceae Justiciaceae

Kadsuraceae Kaliphoraceae Kalmiaceae Kaniaceae Kiggelariaceae Kingdoniaceae Kin

giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae

Krameriaceae

Labiatae Lacandoniaceae Lachenaliaceae Lacistemataceae Lactoridaceae Lactucaceae

Lagerstroemiaceaee LamiaceaeLampsanaceae Lanariaceae Langsdorffiaceae Lantan

aceae Lapageriaceae Lardizabalaceae Lasiopetalaceae LauraceaeLawsoniaceae Laxm

anniaceae Lecythidaceae Ledaceae Ledocarpaceae Leeaceae Leguminosae Leitneriac

eae LemnaceaeLennoaceae Lentibulariaceae Lentiscaceae Leoniaceae Leonticaceae

Lepidobotryaceae LepidocarpaceaeLepidocaryaceae Lepidocerataceae Leptaulaceae

Leptospermaceae Lepturaceae Lepuropetalaceae LeuctenbergiaceaeLibocedraceae

Licaniaceae Ligustraceae Lilaeaceae Liliaceae Limeaceae Limnanthaceae Limnocharit

aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace

ae Linderniaceae Lindleyaceae Linnaeaceae LippayaceaeLiriaceae Liriodendraceae Li

ssocarpaceae Littorellaceae Loasaceae Lobeliaceaee Loganiaceae LomandraceaeLoni

ceraceae Lopeziaceae Lophiocarpaceae Lophiolaceae Lophiraceae Lophophytaceae L

ophopyxidaceae LoranthaceaeLotaceae Lowiaceae Lupulaceaee Luxemburgiaceae Lu

zuriagaceae Lychnidaceae Lyciaceae LyginiaceaeLygodisodeaceae Lysimachiaceae Ly

thraceae

Macarisiaceae Mackinlayaceae Madiaceae Maesaceae Magnoliaceae Malaceae Male

sherbiaceae MalortieaceaeMalpighiaceae Malvaceae Mangiaceae Manicariaceae Ma

paniaceae Marantaceae Marathraceae MarcgraviaceaeMartyniaceae Mastixiaceae

Matricariaceae Maundiaceae Mayacaceae Medeolaceae Medusagynaceae Peridiscac

eaeMelaleucaceae Melampyraceae Melanophyllaceae Melanthiaceae Melastomatac

eae Meliaceae Melianthaceae MelicaceaeMeliosmaceae Melittidaceae Melochiacea

e Memecylaceae Mendonciaceae Menispermaceae Menthaceae MenyanthaceaeMe

nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

Nypaceae Nyssaceae

Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

ceae Papilionaceae Pappophoraceae PapyraceaeParacryphiaceae Parasitaxaceae Par

ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

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hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

49

cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

50

ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 17: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

17

al 2004 Davis et al 2007) it seems likely that Burmanniaceae sl are polyphyletic but

both parts are in Dioscoreales (Merckx et al 2006) and relationships within the largely

hemiparasitic Santalales are also gradually being clarified (Maleacutecot 2002) The inclusion

of parasitic taxa in general molecular analyses can cause conniptions (eg Nickrent et al

2004 Davis et al 2004 Chase et al 2006 G Petersen et al 2006b) and there can be

horizontal transmission of genes (eg Davis amp Wurdack 2005 Vitaceae to Rafflesiaceae

Barkman et al 2007 the mitochondrial atp1 gene commonly moves)

Similarly in water plants neither vessels in particular nor much xylem in general is

needed leaves are highly modified and water-mediated pollination if adopted may

well be associated with major changes in floral morphology Here too recent molecular

studies suggest that aquatic groups with hitherto problematic relationships may find

homes Thus Podostemaceae are close to Clusiaceae and Calophyllaceae and are sister

to Hypericaceae (Malpighiales Kita amp Kato 2001) Hydatellaceae which used to be in

Poales are part of Nymphaeales (Saarela et al 2007) and Hydrostachyaceae may be

close to Hydrangeaceae (Cornales Xiang et al 2002) or perhaps should be placed in

Lamiales note that in the first two cases in particular there are morphological and

chemical features that support such a move If Podostemaceae are indeed close to

Clusiaceae I look forward to seeing hypotheses to explain how the dramatic changes in

the vegetative body that have made Podostemaceae so problematic for generations of

systematists took place That conventional wisdom has trouble in understanding or

explaining how the morphologies of groups like Clusiaceae and Podostemaceae can be

related is largely a problem with conventional wisdom Ceratophyllaceae seem to be

finding a position as sister to eudicots (see Moore et al 2007) but their morphology is

so derived that there is no morphological evidence of which I am aware for this

relationship There are also similar although less extreme morphological problems with

many plants pollinated by wind

In such situations especially with some water plants mycoheterotrophs and

parasites a variety of characters may be affected and particularly under such

circumstances groups have tended to be formed using a few or even only a single

character that seems to provide evidence of relationships Add to this the tendency to

weight some characters particularly strongly on a priori grounds and the result was

either the recognition of conglomerate taxa such as Amentiferae which now appear to

be highly polyphyletic or the segregation of families like Plantaginaceae s str (now

much expanded) and Leitneriaceae (now in Simaroubaceae) - all of these are more or

less wind pollinated

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

18

Even although relationships of many parasitic and aquatic groups are being

clarified and at least some of the proposed phylogenetic positions seem to be well

supported it is indeed often still very difficult to understand how such groups have

evolved from their more morphologically conventional relatives However we have had

all sorts of preconceptions as to how characters should evolve Thus we used to

assume that features like highly scalariform vessel perforation plates or the complete

absence of vessels or a flower with an androecium that had many stamens a superior

ovary or separate petals were necessarily plesiomorphic or primitive and conversely

a simple perforation plate and the presence of vessels an androecium with few

stamens inferior ovary or petals that were connnate were almost necessarily

apomorphic or advanced (but cf eg Stebbins 1951) Such assumptions are incorrect

(eg Soltis et al 2005b) Carpels may become secondarily free carpels may fail to close

the seeds then developing outside the confines of the carpel as in some Aspagaraceae-

Nolinoideae Violaceae Berberidaceae Malvaceae-Sterculioideae etc) in Peliosanthes

teta perhaps the only species in Peliosanthes(Asparagaceae-Nolinoideae) the ovary

varies from superior to inferior (Jessop 1976 species limits here need close

investigation see also Kuzoff et al 2001 and Soltis amp Hufford 2002 Saxifragaceae

Apiales etc) many-seeded carpels can evolve from few-seeded carpels

(Razafimandimbison et al 2008) monoecy may be derived from dioecy (Schaefer amp

Renner 2010 and references) Classic studies such as those by Babcock (eg 1947)

on Crepis that assumed that evolution - in this case of the karyotype in particular - was

unidirectional have needed comprehensive re-evaluation (Enke amp Gemeinholzer 2008)

Most if not all characters have reversed andor evolved in parallel as is clear in the

discussion of ovary position here especially in Poales Saxifragales and Asparagales and

also in comments on such morphologically distinctive taxa (when compared with their

immediate relatives) such as Menyanthaceae and especially Pittosporaceae Parallel

evolution may occur even at the level of amino acid substitution as in the independent

acquisition of the phosphoenolpyruvate carboxylase (pepC) gene in C4 photosynthesis in

grasses (Christin et al 2007b see also Blaumlsing et al 2000)

Thinking of how such characters have been used in the past in classifications at the

generic level the results have been similar to those just discussed above for parasitic

plants and others Relying too much on animal pollination syndromes to mark generic

boundaries has all too often led to taxa that are highly unsatisfactory phylogenetically

(see eg Acanthaceae Bignoniaceae Campanulaceae Ericaceae Lamiaceae

Orchidaceae) and over-reliance on characters of fruit and seed (see eg Brassicaceae

and Apiaceae) has also led to unsatisfactory generic limits Again the more general

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

19

problem is the use of one or a very few characters that have been weighted a priori to

structure classifications (see also Garciacutea et al 2009) although it is perhaps not

surprising that such an approach sometimes results in the delineation of taxa that have

indeed turned out to be quite distinct (cf eg Burtt Astragalus versus Oxytropis)

Taxa recognized here are monophyletic yet many are polythetic at the

morphological level that is they lack unique features characterizing (defining) all and

only members of that family they can be recognised phenetically only by the unique

combinations of characters that they posess This is the result of evolution any

synapomorphy characterizing a taxon may be lost or modified beyond easy recognition

in some of its members or the synapomorphy may appear to be identical to a feature

that has evolved in parallel in a quite unrelated plant That plant groups are polythetic is

almost as much a feature of monophyletic taxa as the paraphyletic taxa common in

evolutionary classifications Some families now include substantial variation as

phenetically distinct derived groups are placed in their proper phylogenetic position -

examples are Ericaceae which include the erstwhile Empetraceae a wind-pollinated

group that is florally very different and the overwhelmingly large-flowered and animal-

pollinated Plantaginaceae which now include several derived small-flowered aquatic

and wind-pollinated groups that were previously placed in separate families

Thus some families as delimited here may not be easy to recognize However

remember that detecting relationships - use whatever characters you can even if they

are not obvious - and naming a plant - focus on easy-to-see characters that may not

reflect relationships - are quite different problems Taxa although natural may not be

readily recognizable indeed it was in exactly this context that Lamarck worked out the

basic principles of writing dichotomous keys in 1778 (see Scharf 2007 for keys and the

like) Of course Lamarcks idea of nature was very different from ours - he thought that

there was some kind of continuum of form on which living organisms were to be

situated with no real gaps anywhere - but this meant that his genera (for example)

might well not be sharply distinct from each other even if each were part of the real

continuum that was life Lamarcks separation of the establishment of relationships and

groupings in a system and identification of the plants in that system have been widely

accepted (but cf eg Godfray amp Knapp 2004 many of the contributors to Heywood et

al 2007) Perhaps the best way of identifying plants at the family level is by well-made

multiple access keys as in Watson and Dallwitz (1992a onwards family limits there may

differ substantially from those adopted here) Multiple access keys free users from the

constraints of dichotomous keys in which particular characters are needed at each step

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

20

of the identification process before there can be further progress Instead those

characters that happen to be evident on a specimen can be used in whatever order is

convenient when linked to illustrations glossaries etc their power is enormous (see

Dallwitz et al 2000 [2006] for the principles underlying their construction and use)

Nevertheless dichotomous keys such as those of Hutchinson (1973) and Franz Thonner

(Geesink et al 1981) have their uses Of course taxa coming out adjacent in keys may

well not be at all related

(When identifying large numbers of plants even more efficient than either style of

identification and certainly lots more fun is sight identification Unless you have a

photographic memory you have to build up your knowledge of comparative plant

morphology - on which the ability to make accurate identifications depends - by

repeated observation When faced with an unknown plant I always look for leaf teeth

and stipules distinctive hair types especially stellate and T-shaped hairs glands and

punctations of any sort the presence of latex or other exudate and check leaf insertion

smelling crushed leaves can also be helpful In this context nodal anatomy can usually

be checked using a razor and a hand lens or even simply carefully examining leaf scars

The short paragraphs added after most families may help in confirming familial

identifications)

For the record and for the little that it is worth there are 4 orders and 13 families

of gymnosperms characterised on these pages and together they include some 82

genera and 947 species For angiosperms comparable figures are 56 orders 445

families 13208 genera and 261750 species (of which monocots include 11 orders 89

families 2759 genera and 52760 species) Note however that higher mathematics

was never my strong point and anyway these are pretty meaningless figures even for

species which many (but not all) might concede smacked slightly more of reality than

other taxa estimates range as high as 422000 (Govaerts 2001) Furthermore numbers

of genera and species change daily Nevertheless as emphasized here families are

useful in teaching we as a community can ensure that their limits remain largely stable

and by concentrating on relatively few of them one can gain some familiarity with much

of the worlds flora For summaries of the sizes of orders and families and of the general

arrangement followed in these pages see also the Statistics page itself

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

21

A PHYLOGENETIC TREE OF SEED PLANTS (THE MAIN TREE)

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

22

The Main Tree above is a conservative summary of well supported relationships

between the major clades of extant seed plants Reading - interpreting and

understanding - such a phylogenetic tree is not necessarily easy so reading a tree may

require a bit of practice OHara (eg 1988 1992) provides an accessible introduction to

thinking and talking about history in the context of looking at phylogenetic trees More

recently Krell and Cranston (2004) Crisp and Cook (2005) and others have emphasized

how careful one must be when interpreting and talking about evolution in ladderized

trees in particular and phylogenetic trees in general Gregory (2008) provides a

particularly detailed discussion of this problem In ladderized trees the smaller (in terms

of numbers of terminals) sister taxon is consistently shown on the same side at every

node thus the trees tend to be pectinate like the teeth of a comb however the

horizontal axis that is as a result so evident has no polarity or particular direction The

use of the adjective basal when discussing phylogenetic trees is especially dangerous

(see also Wojciechowski et al 2004 D Soltis et al 2005b) When I use the term and the

context is not otherwise clear I am referring to the pectinations at the base of a

ladderised tree Remember that when talking about sister taxa one can never be basal

to the other or older or younger although members of one can be more derived - in the

sense of having more apomorphies - than the other (but even then one has to be

careful) The words primitive and advanced and lower and higher should

practically never be used especially when talking about taxa Amborellaceae and

Pinaceae are sister to all other flowering plants and Pinales and are basal to the crown

groups of all other flowering plantsPinales respectively but that does not mean they

are thereby primitive Pinaceae in particular have numerous apomorphies The word

plesiomorphic is far less loaded than primitive and can be used to talk about

individual characters

Most trees here have been more or less ladderized as is the one above The

ladderization may be imperfect for example you can see that on the Main Tree

asterids with ten orders follow rosids and relatives which have seventeen although

this is in part because of the currently unresolved nature of relationships between the

main clades of core eudicots (this is likely to change soon) The Main Tree could have

been drawn with Amborella Acorus or a host of other taxa at the far right without

offending any relationships phylogenetic trees are like mobiles the only fixed points

being the nodes Despite the problems just mentioned pectination interpreted

carefully has its value and it is not an arbitrary process As one reads the terminals of a

pectinate tree from left to right adjacent terminals will be separated by apomorphies

that will cumulate in an appropriate fashion When thinking of a book or a herbarium

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

23

sequence (see Haston et al 2007) this is of some value Nymphaeales and

Austrobaileyales are here adjacent on the tree but they could be separated by

hundreds of families in the sequence merely by rotating only the node from which

Nymphaeales and their sister group (all other angiosperms minus about 75 species)

arise If adjacent in a book or herbarium or as in these summary pages then it is

relatively easy to relate their characters both apomorphic and plesiomorphic but if

separated by hundreds of pages or two floors in a large building then it is less easy to

get anything from the sequence (Since all orders in the pages of the main part of this

site are preceded by the apomorphies of all nodes immediately below them in the seed

plant phylogeny and because of the linkages that have been built in to the site this

problem is much less serious there there is no sequence) So the particular sequence of

all seed plants that is based on a phylogenetic classification but is to be used in herbaria

or any other place where taxa have to be arranged linearly as in this part of APweb

will be one that maximizes the number of taxa that are both successive branches of the

tree and placed successively in the sequence Since specimens are generally filed under

families the outline of a new family sequence for arranging herbaria and books can now

be suggested (Haston et al 2007) and such a standardized sequence will further help

teaching and learning about plants

ON THINKING ABOUT APOMORPHIES

Identifying apomorphies is important because understanding the evolution of

morphology in the broad sense - ie including anatomy chemistry etc - in the context

of diversification is one of our major goals For identifying apomorphies several

preconditions must be met One needs to have an accurate well supported phylogeny

one has to have examined the right taxa both from the point of view of morphology and

molecules one has to have coded the characters right (ie delimited states

appropriately) and one has to use an appropriate model of evolution when fitting the

variation to the tree (see eg Omland 1999 Stevens 2006b) If the distributions of

apomorphies below are compared with those in other studies (eg Turgeon et al 2001

Bremer et al 2001 Endress 2001 Albach et al 2001a Judd amp Olmstead 2004 D Soltis

et al 2005b Zhang et al 2006 Judd et al 2007) the reader may find a number of

differences Although I have integrated such studies as far as is possible there are five

reasons that link to the preconditions just mentioned why there may be differences

where features are placed on trees

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

24

1 I may not have found all the information about a particular character there may be

disagreement over its interpretation or I have added information

2 The sampling of nearly all molecular studies is very incomplete (see Salisbury amp Kim

2001 for problems caused by sampling) indeed it is commonplace to decry the

incompleteness of molecular sampling But not only is the sampling in molecular studies

often less than we might wish that of the morphological and chemical characters whose

evolution we are interested in understanding is also often very poor So for many

anatomical chemical and embryological characters that are confidently said to

characterise families and other groups we all too often have no idea if those characters

are applicable to the whole clade or just to a subgroup within that clade However as

our sampling improves we can locate changes on the tree more precisely if in

unexpected positions Thus is is sometimes suggested that Ericaceae have ellagic acid

(Soltis et al 2005b) in fact the little evidence we have indicates that only a small clade

of some 80 species has ellagic acid and the rest of the family - itself alone about one

third of all Ericales - does not Similarly Ericaceae are noted for their

ectendomycorrhizal associations but Enkianthus sister to all other Ericaceae

apparently lacks these (Abe 2005) Thus these mycorrhizae and their associated

distinctive hair roots are unlikely to be apomorphies of Ericaceae - however it appears

that other families near Ericaceae may also have ectendomycorrhizae (see Asai 1934)

3 I am fitting characters to a very conservative tree with many polytomies although the

nodes that are utilised are for the most part strongly supported polytomies make the

optimisation of characters that is the assigment of character state change to a

particular node on the tree notably difficult (eg Madison amp Madison 2002) In nearly

all studies of the evolution of characters distributions of characters are optimised on a

more or less fully resolved tree and the construction of supertrees may yield yet more

detailed hypotheses of relationships (for literature on supertrees see Cotton amp

Wilkinson 2007 2008) Of course some nodes on such fully resolved trees andor

supertrees may have little support and optimisations of characters on such trees may

carry correspondingly little conviction Even parts of some of the trees used here have

poor support eg relationships within aquatic Alismatales etc although I indicate

when this is the case

4 Exactly how one goes about optimising a character on a tree is critically important

Even using simple parsimony optimisations (ACCTRAN or DELTRAN ACCelerated

TRANsitions or DELayed TRANsitions) the position of synapomorphies on trees - and

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

25

hence our ideas of evolution - may differ and this is still more true if one uses maximum

likelihood or Bayesian methods and makes apparently reasonable suggestions about

weighting gains over losses (eg Donoghue amp Ackerley 1996 Cunningham et al 1998

Omland 1997 1999 Ree amp Donoghue 1999 Polly 2001 Webster amp Purvis 2001

Ronquist 2004 Crisp amp Cook 2005) Sannier et al (2007) show how in in Arecaceae that

where on a phylogenetic tree one might peg changes in microsporogenesis will depend

on the methods one uses to do this and Pedersen et al (2007) discuss the sometimes

very substantial effect of node support on the posterior probabilities of ancestral

character states Here I use parsimony optimization not always as explicit as it might be

but I have often indicated where there are particularly important uncertainties as to the

positions of particular character changes on the tree

5 Finally although I have paid quite a lot of attention to the delimitation of the

character states that make up all the characterizations I have not spent enough time on

this critical operation If we are interested in understanding evolution then fitting the

basic variation - not character states - to a tree in principle allows greater flexibility in

understanding morphology in the context of local phylogenies (see also Stevens 2000

Endress 2005c) However many character states used here are delimited globally that

is they are circumscribed in the context of the variation shown by individual characters

across all angiosperms andor in the context of classic ideas of character evolution

Character states often have arbitrary limits and serve best to communicate

information whether they are in fact suitable for either phylogenetic analysis or

understanding evolution are separate issues Studies have rather unsurprisingly

perhaps but importantly shown that dividing the one character into different sets of

states may yield differing ideas of evolution of that character (eg Lamb Frye amp Kron

2003 Hibbett 2004) When looking at trees on which character states are optimised

one should bear in mind the problems surrounding the delimitation of states (eg

Stevens 2000 2006b) and the danger of using pollen or other types - constructs

based on many characters that vary independently but which effectively get lost in

these types thus Blackmore et al (2009) decomposed the pollen types in Asteraceae

into 52 characters

I use Remanes three main criteria of homology or better similarity when

determining the basic similarity of structures on different organisms (see Remane 1952)

These criteria are special properties position and intermediates Special properties

include anatomical or chemical characters gene expression data etc Position refers

to the position of an organ with respect to landmarks on the plant Although plants are

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

26

plants and landmarks are not as fixed as one might like trying to work out the relative

positions of parts is a good way of understanding morphology so long as one does not

try to out-do Procrustes Intermediates include intermediates found on the same

plant or on different plants Intermediates may observed during development or be

adult structures Thus one may find changes during development which suggest what

the nature of a particular structure is and very different stuctures on different plants

may turn out to be similar early in development Or one can simply compare different

taxa and see that structures that appear to be very different when comparing two

different taxa can be linked morphologically by looking at other taxa

Nevertheless the use of these criteria may not yield an unambiguous answer as to

what a structure is even given a solid phylogeny and an improved understanding of

development (see Jaramillo amp Kramer 2007 for a useful discussion) As Endress (2005c)

observed a number of features - position function development shape anatomy

histology gene activity and relationships to other taxa that clearly have petals - can be

used to distinguish a petal (for example) from other floral structures if a petal does not

have one of these features is it thereby not a petal Thus Maturen et al (2005) recently

found that floral organ diversity genes (B and C) were expressed in the large white

inflorescence bracts of Cornus (see also Costa et al 2005) Peney et al (2005) noted that

not all monosulcate pollen grains in monocots have the same developmental pathway

and that as a result such pollen might not have the same ancestral state Reeves and

Olmstead (2003) suggested that the genetic mechanisms causing monosymmetry in

Lamiales and Solanales were different and Serna and Martin (2006) described similar

problems with the development of hairs in Arabidopsis when compared with that of

hairs in Antirrhinum and Solanaceae Indeed as one perhaps might expect delimitation

of states and characters does not necessarily become easier with increasing knowledge

of development etc Thus Buzgo et al (2004) Matthews and Endress (2005) and others

have shown how hard it can be to distinguish between eg prophylls and other floral

structures as their behaviour is studied during the course of the development of the

flower Similarly at what concentration is a particular secondary metabolite deemed to

be present (Waterman (2007)

The validity of the approach used here that of fitting morphological variation to a

largely molecular-based tree may be questioned However I think it rather unlikely that

well-supported molecular branches will be overturned by morphological data Indeed

analyses of morphological data alone do provide support for many of the clades evident

in molecule-only analyses and analyses with morphological and molecular data

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

27

together may lead to increased support for clades (eg Hufford 1992 Nandi et al 1998

[but adding morphological data reduces support for a number of critical clades too]

Doyle amp Endress 2000 however in none of these papers is the use of morphology

without ambiguity) It is unfortunately clear that the use of morphology alone may not

suggest problems in the phylogenetic placement of taxa that later turn out to have been

wrongly included (eg Zhang et al 1992) Thus I have been wary of putting much weight

on clades that have only morphological support but note that I have treated molecule-

based clades with low bootstrap or jacknife support values (esp below 70) or low

posterior probabilies (below 095) likewise Although I may have been mistaken in

placing so much emphasis on molecular data in terms of providing the basic

phylogenetic framework for angiosperms morphological and molecular data are only

very rarely in irreconcilably strong conflict There are indeed a few places where the

conflict seems extreme These include the relative positions of the Monimiaceae and

Hernandiaceae (Laurales) the position of Hanguanaceae (Commelinales [as here] or

Zingiberales) and of Triplostegia (is it in Dipsacaceae or Valerianaceae - see

Dipsacales) Fossils are unlikely to affect the topologies of the trees presented here but

see below for their importance in understanding morphological evolution in general and

the evolution of angiosperms in particular Nevertheless some largely reject the idea

that trees based on molecular data alone can recover phylogenetic relationships

especially when branching points are old and prefer to used trees based on analysis of

morphological data including those taken from fossils (Hilton amp Bateman 2006 Farjon

2007)

All in all however the extent of the congruence between morphological and

molecular data is impressive and heartening and many clades can be characterised

morphologically It seemed in 1998 that there were no unambiguous morphological

synapomorphies for angiosperm orders (K Bremer 2000) and this is still true if by

unambiguous is meant non-homoplasious However many orders and other clades

have synapomorphies even if these may be indistinguishable at least at the current

level of morphological and developmental knowledge from parallel occurences

elsewhere As our morphological knowledge increases so too does the number of

apomorphies

Endress and Matthews (2006a) emphasize the importance of tendencies and

developmental constraints when thinking about characters of clades within the rosids -

many characters or character combinations occur in a rather sporadic fashion within

the clade and apparently notably less frequently outside it There are a number of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

28

examples of tendencies here and in the individual order pages such as the features

enclosed by parentheses in groups above the level of family eg see (cuticular waxes

as aggregated rodlets) for the commelinids and the discussion of the distribution of

polyandry that is flowers with numerous stamens in the asterid I + II groups etc

Tendencies may involve more than single characters When thinking of the

associations of features like integument thickness and vascularization of the integument

(Endress amp Matthews 2006a) one can imagine fairly simple developmental

preconditions being operative As Endress and Matthews (2006a) note it is difficult to

imagine vascular tissue developing in an integument only two cells thick Changes here

would be loosely correlated if morphologically linked However the strongly correlated

changes noted by Givnish et al (2005) are ecologically linked but are presumably

morphologicallydevelopmentally independent When there seem to be characters

evolving more or less together tests can be carried out to see if the changes are

concentrated on certain branches of the tree (eg Maddison 1990 Sanderson 1991

Maddison amp Maddison 2000) Such tests have rarely been carried out at higher levels in

angiosperms

One of the most striking examples of a tendency is the distribution of N-fixation

restricted as it is to a monophyletic group of four clades although it has arisen seven (or

perhaps several more) times independently within the N-fixing clade and several

members of two quite different kinds of bacteria are involved (eg Clawson et al 2004

Elliott et al 2007 Sprent amp James 2007) Other examples of work that bears on the issue

of tendencies include the findings that flowers of polysymmetrical Arabidopsis have

genes like TCP1 that are expressed asymmetrically during early development and TCP1

is a probable orthologue of the well-known CYC gene of Antirrhinum that is involved in

the development of monosymmetric slowers there (Cubas et al 2001 Costa et al 2005

etc) Parallelism might build on this underlying morphologically cryptic monosymmetry

even if details of the genetic mechanisms causing the monosymmetry evident in

particular groups may be different (Reeves amp Olmstead 2003 Cubas 2004 see above)

Similarly there have been several recent suggestions that the capability to synthesise a

particular metabolite may be switched off but not lost and so can sometimes be

reacquired (eg Wink amp Witte 1983 Wink 2003 Liscombe et al 2005 Larsson 2007

Waterman 2007) Hence perhaps the rather spotty distribution of many secondary

metabolites like ellagic acid the indole alkaloid camptothecin iridoids etc when

considered in the context of phylogenies As a non-botanical example - but a rather nice

one - Salwini-Plawen and Mayr (1961) suggested some time ago that there has been

considerable parallelism (40-65 or more independent origins) in the evolution of eyes in

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

29

metazoans However Pax 6 seems to be a master control gene that is involved in eye

formation perhaps throughout all bilateralians (eg Gehring amp Ikeo 1999 Erwin amp

Davidson 2002) which changes the nature of the problem Note however that in

general our invocation of evolutionary tendencies to explain the patterns of

distributions of characters is really an admission that we do not understand those

patterns

As we find out more about variation we will probably find fewer and fewer features

constant throughout a group Most unqualified statements of presence and absence

should properly be qualified as usually present or usually absent if one is thinking of

the characterisations on the order pages as encompassing the total variation within a

clade Thus Pistia alone among monocots as so far known has sieve tube plastids with

starch grains not protein crystals However this does not affect the fact that sieve tube

plastids with cuneate protein crystals and lacking starch grains are an apomorphy for

monocots Furthermore fossils in a number of cases suggest character combinations

unknown in extant taxa as may be seen in the discussions of Fagaceae Platanaceae

Iteaceae Calycanthaceae etc Confusing the issue there may be questions as to where

exactly on the tree a particular fossil is to be placed (eg see Nymphaeaceae

Calycanthaceae Archaefructus etc)

To summarize given our current understandings of both phylogenies and

characters evolution of some characters in which we are interested seems very labile

(see eg D Soltis et al 2005b Endress and Matthews 2006a Stevens 2006b Ekman et

al 2008 for a good recent study) and I have been cautious when talking about character

evolution Much effort must continue to be spent in summarizing characters of well-

established clades at all levels providing features by which they may be recognized and

signaling synapomorphies Remember that (1) the basic morphological anatomical and

chemical knowledge of many critical taxa is woefully incomplete (2) different

assumptions about character evolution may greatly affect the position of

synapomorphies on trees (3) in many cases relationships within and between many

groups are too uncertain at present to worry very much about synapomorphies and

(4) we must be clear about what we do and do not not know As mentioned above I

have indicated in a number of places where there are particularly important

uncertainties as to where characters should be placed on the tree Nevertheless it is a

relatively easy matter to update notes such as these and it can be a simple matter to

incorporate new data on characters that have never before been considered in the

context of a tree Much basic - and unfortunately perhaps unfashionable - work must be

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

30

carried to clarify the distribution of morphological anatomical and chemical characters

for examples of what can to be done see the work of P K Endress and collaborators

(floral morphology and development) and S R Jensen and collaborators (iridoids) But

acquiring information about nodal anatomy cell and tissue distribution nodal anatomy

and the like is not difficult at all and should be generally encouraged

But all this is merely a necesary prelude to the understanding of evolution For this

one needs to know a lot more including dating the phylogeny understanding the

function(s) of characters factoring in relevant aspects of the palaeoenvironment in

which an apomorphy first appeared etc

SUMMARY OF THE SYSTEM

Below is a formalised summary of the relationships within orders of the families of seed

plants There are a few families that are not recognised even as options in APG II and

vice versa as well as a few extra orders The families to be recognised in the most

recent edition of Mabberleys The Plant Book (Mabberley 2008) are also largely

consistent with those below But all differences are trivial and will - I hope - eventually

disappear

Square brackets - [] - enclose clades the plus sign - + - designates sister taxa a

comma - - denotes part of a polytomy and quotation marks - - denotes a

paraphyletic group

SEED PLANTS

GYMNOSPERMS

Cycadales

Cycadaceae + Zamiaceae

Ginkgoales

Ginkgoaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

31

Pinales

Pinaceae [[Araucariaceae [Podocarpaceae + Phyllocladaceae]] [Sciadopityaceae

[Taxaceae + Cupressaceae]]]

Gnetales

Ephedraceae [Gnetaceae + Welwitschiaceae]

ANGIOSPERMSFLOWERING PLANTS

Amborellales

Amborellaceae

Nymphaeales

Hydatellaceae [Cabombaceae + Nymphaeaceae]

Austrobaileyales

Austrobaileyaceae [Schisandraceae + Trimeniaceae]

Chloranthales

Chloranthaceae

MAGNOLIIDS

Magnoliales

Myristicaceae [Magnoliaceae [[Himantandraceae + Degeneriaceae] [Eupomatiaceae +

Annonaceae]]]

Laurales

Calycanthaceae [[Siparunaceae [Gomortegaceae + Atherospermataceae]] [Monimiaceae

[Hernandiaceae + Lauraceae]]]

Canellales

Canellaceae + Winteraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

32

Piperales

[Hydnoraceae + Aristolochiaceae] [Piperaceae + Saururaceae]

MONOCOTS

Acorales

Acoraceae

Alismatales

Araceae [Tofieldiaceae [[Alismataceae [Hydrocharitaceae + Butomaceae]]

[Scheuchzeriaceae [Aponogetonaceae [Juncaginaceae [Maundiaceae [[Posidoniaceae

[Ruppiaceae + Cymodoceaceae]] [Zosteraceae + Potamogetonaceae]]]]]]]

Petrosaviales

Petrosaviaceae

Dioscoreales

Nartheciaceae [[Taccaceae + Thismiaceae] [Burmanniaceae + Dioscoreaceae]]

Pandanales

Velloziaceae Triuridaceae Stemonaceae [Pandanaceae + Cyclanthaceae]

Liliales

Corsiaceae [Campynemataceae [Petermanniaceae [Colchicaceae + Alstroemeriaceae]]

Melanthiaceae [[Philesiaceae + Rhipogonaceae] [Smilacaceae + Liliaceae]]]

Asparagales

Orchidaceae [[Boryaceae [Blandfordiaceae [Lanariaceae [Asteliaceae + Hypoxidaceae]]]]

[[Ixioliriaceae + Tecophilaeaceae] [Doryanthaceae [Iridaceae [Xeronemataceae

[Xanthorrhoeaceae [Alliaceae + Asparagaceae]]]]]]]

COMMELINIDS

Unplaced

Dasypogonaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

33

Arecales

Arecaceae

Poales

[Typhaceae + Bromeliaceae] [Rapateaceae [[Thurniaceae [Juncaceae + Cyperaceae]]

[[Mayacaceae [Eriocaulaceae + Xyridaceae]] [Flagellariaceae [Anarthriaceae

[Centrolepidaceae + Restionaceae]]] [Joinvilleaceae [Ecdeiocoleaceae + Poaceae]]]]]]]

Commelinales

[Commelinaceae + Hanguanaceae] [Philydraceae [Haemodoraceae + Pontederiaceae]]

Zingiberales

Musaceae [Strelitziaceae + Lowiaceae] Heliconiaceae [[Cannaceae + Marantaceae]

[Costaceae + Zingiberaceae]]

Ceratophyllales

Ceratophyllaceae

EUDICOTS

Ranunculales

Eupteleaceae [Papaveraceae [[[Lardizabalaceae + Circaeasteraceae] [Menispermaceae

[Berberidaceae + Ranunculaceae]]]]

Sabiales

Sabiaceae

Proteales

Nelumbonaceae [Platanaceae + Proteaceae]

Trochodendrales

Trochodendraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

34

Buxales

Haptanthaceae [Buxaceae + Didymelaceae]

CORE EUDICOTS

Gunnerales

Gunneraceae + Myrothamnaceae

Dilleniales

Dilleniaceae

Saxifragales

Peridiscaceae [[Paeoniaceae [Altingiaceae [Hamamelidaceae [Cercidiphyllaceae +

Daphniphyllaceae]]]] [[Crassulaceae [Aphanopetalaceae [Tetracarpaeaceae

[Penthoraceae + Haloragaceae]]]] [Iteaceae [Grossulariaceae + Saxifragaceae]]]]

Cynomoriaceae unplaced

Vitales

Vitaceae

ROSIDS

FABIDROSID I

Zygophyllales

Krameriaceae + Zygophyllaceae

Celastrales

Lepidobotryaceae + Celastraceae

Oxalidales

Huaceae [[Connaraceae + Oxalidaceae] [Cunoniaceae [Elaeocarpaceae [Brunelliaceae +

Cephalotaceae]]]]

Malpighiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

35

[Achariaceae [Goupiaceae [Violaceae + Passifloraceae] [Lacistemataceae Salicaceae]]

[Lophopyxidaceae Putranjivaceae] [Ctenolophonaceae [Erythroxylaceae +

Rhizophoraceae]] Linaceae Ixonanthaceae Humiriaceae Irvingiaceae

Centroplacaceae Pandaceae [[Bonnetiaceae + Clusiaceae] [Calophyllaceae

[Hypericaceae + Podostemaceae]]] [Malpighiaceae + Elatinaceae] Ochnaceae

[Peraceae [Rafflesiaceae + Euphorbiaceae]] [Phyllanthaceae + Picrodendraceae]

[Balanopaceae [[Trigoniaceae + Dichapetalaceae] [Chrysobalanaceae +

Euphroniaceae]]] Caryocaraceae

N-FIXING CLADE

Fabales

Quillajaceae [Fabaceae [Polygalaceae + Surianaceae]]

Rosales

Rosaceae [[Barbeyaceae [Elaeagnaceae Dirachmaceae Rhamnaceae]] [Ulmaceae

[Cannabaceae [Moraceae + Urticaceae]]]]

Cucurbitales

Anisophylleaceae [[Corynocarpaceae + Coriariaceae] [Cucurbitaceae [Tetramelaceae

[Datiscaceae + Begoniaceae]]]] Apodanthaceae

Fagales

Nothofagaceae [Fagaceae [[Myricaceae + Juglandaceae] [Casuarinaceae

[Ticodendraceae + Betulaceae]]]]

MALVIDROSID II

pgtGeraniales

Geraniaceae [[Melianthaceae + Francoaceae] [Vivianaceae + Ledocarpaceae]]

Myrtales

Combretaceae [[Onagraceae + Lythraceae] [[Vochysiaceae + Myrtaceae]

[Melastomataceae [Crypteroniaceae [Alzateaceae + Penaeaceae]]]]]

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

36

Crossosomatales

[Staphyleaceae [Guamatelaceae [Crossosomataceae + Stachyuraceae]]] [Aphloiaceae

[Geissolomataceae + Strasburgeriaceae]]

Picramniales

Picramniaceae

Sapindales

Biebersteiniaceae Nitrariaceae [Kirkiaceae [Anacardiaceae + Burseraceae]]

[Sapindaceae [Simaroubaceae Rutaceae Meliaceae]]

Huerteales

Gerradinaceae [Dipentodontaceae + Tapisciaceae]

Malvales

Neuradaceae [Thymelaeaceae [Sphaerosepalaceae Bixaceae [Cistaceae

[Sarcolaenaceae + Dipterocarpaceae]] [Cytinaceae + Muntingiaceae] Malvaceae]]

Brassicales

[Akaniaceae + Tropaeolaceae] [[Moringaceae + Caricaceae] [Setchellanthaceae

[Limnanthaceae [[Koeberliniaceae [Bataceae + Salvadoraceae]] [Emblingiaceae

[Pentadiplandraceae [Gyrostemonaceae Resedaceae Stixis and relatives] Tovariaceae

[Capparaceae [Cleomaceae + Brassicaceae]]]]]]]]

Berberidopsidales

Aextoxicaceae + Berberidopsidaceae

Santalales

Erythropalaceae Olacaceae [[Loranthaceae [Misodendraceae + Schoepfiaceae]]

[Opiliaceae + Santalaceae]] Balanophoraceae unplaced

Caryophyllales

[[Droseraceae [Nepenthaceae [Drosophyllaceae [Ancistrocladaceae +

Dioncophylleaceae]]]]] [[Frankeniaceae + Tamaricaceae] [Polygonaceae +

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

37

Plumbaginaceae]]] [Rhabdodendraceae [Simmondsiaceae [[Asteropeiaceae +

Physenaceae] [[Caryophyllaceae [Achatocarpaceae + Amaranthaceae]]

[Stegnospermataceae [Limeaceae [[Lophocarpaceae [Barbeuiaceae [Aizoaceae

[Sarcobataceae Phytolaccaceae Nyctaginaceae]]]] [Molluginaceae [Halophytaceae

Montiaceae Basellaceae Didiereaceae [Talinaceae [Portulacaceae [Anacampseros etc +

Cactaceae]]]]]]]]]]]]]

ASTERIDS

Cornales

[Cornaceae + Nyssaceae] [Hydrangeaceae + Loasaceae] Hydrostachyaceae [Curtisiaceae

+ Grubbiaceae]

Ericales

[Balsaminaceae [Marcgraviaceae + Tetrameristaceae]] [[Polemoniaceae +

Fouquieraceae] Lecythidaceae [[Sladeniaceae + Pentaphylacaceae] [Sapotaceae

[Ebenaceae [Maesaceae [Theophrastaceae [Myrsinaceae + Primulaceae]]]]]

[Mitrastemonaceae Theaceae [Symplocaceae [Styracaceae + Diapensiaceae]]

[[Sarraceniaceae [Roridulaceae + Actinidiaceae] [Clethraceae [Cyrillaceae +

Ericaceae]]]]]]

LAMIIDASTERID I

Unplaced

Metteniusaceae Oncothecaceae Icacinaceae and some unplaced ex-Icacinaceae

Garryales

Garryaceae + Eucommiaceae

Unplaced

Boraginaceae Vahliaceae

Gentianales

Rubiaceae [Gentianaceae [Loganiaceae [Gelsemiaceae + Apocynaceae]]]

Lamiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

38

Plocospermataceae [Oleaceae [Tetrachondraceae [Calceolariaceae

[Gesneriaceae Peltanthera [Pedaliaceae Carlemanniaceae Martyniaceae Stilbaceae

Plantaginaceae Linderniaceae Bignoniaceae Verbenaceae Lamiaceae Paulowniaceae

Schlegeliaceae Thomandersiaceae Phrymaceae Rehmannia Acanthaceae

Scrophulariaceae Orobanchaceae Byblidaceae Lentibulariaceae]]]]]

Solanales

[Montiniaceae [Sphenocleaceae + Hydroleaceae]] [Convolvulaceae + Solanaceae]

CAMPANULIDASTERID II

Aquifoliales

[[Cardiopteridaceae + Stemonuraceae] [Aquifoliaceae [Helwingiaceae +

Phyllonomaceae]]

Asterales

[Rousseaceae + Campanulaceae] Pentaphragmataceae [[Alseuosmiaceae [Phellinaceae

+ Argophyllaceae]] Stylidiaceae [Menyanthaceae [Goodeniaceae [Calyceraceae +

Asteraceae]]]]

Escalloniales

Escalloniaceae

Bruniales

[Bruniaceae + Columelliaceae]

Apiales

Pennantiaceae [Torricelliaceae [Griseliniaceae [Pittosporaceae [Araliaceae

[Myodocarpaceae + Apiaceae]]]]]

Paracryphiales

Paracryphiaceae

Dipsacales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

39

Adoxaceae [Diervillaceae [Caprifoliaceae [Linnaeaceae [Morinaceae [Dipsacaceae +

Valerianaceae]]]]]

LINKS TO ORDERS AND FAMILIES

[Back to Top]

MAIN GROUPINGS

Asterids commelinids core eudicots asterid 1 asterid

2 eudicots gymnosperms Magnoliophyta monocots N-fixing claderosids seed plants

ALPHABETICAL LISTING OF ALL ORDINAL NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | L | M | N | O | P | Q | R | S | T | U | V | W | X | Z

Abietales Acanthales Acerales Acorales Actinidiales Actinostrobales Adoxales Aescul

ales Agavales Aizoales AkanialesAlismatales Alliales Alseuosmiales Alstroemeriales

Altingiales Amaranthales Amaryllidales Amborellales AmbrosialesAmmiales Amomal

es Ancistrocladales Anisophylleales Annonales Anthobolales Apiales Apocynales Apo

nogetonalesAquifoliales Arales Araliales Aralidiales Araucariales Arecales Aristolochi

ales Athrotaxidales Asarales AsclepiadalesAsparagales Asphodelales Asparagales Ast

eliales Atriplicales Aucubales Austrobaileyales Avenales

Balanitales Balanopales Balanophorales Balsaminales Barbeyales Barclayales Batales

Begoniales BerberidalesBerberidopsidales Betulales Biebersteiniales Bignoniales Bix

ales Boraginales Brassicales Brexiales Bromeliales BrunialesBrunoniales Burmannial

es Burserales Butomales Buxales Byblidales

Cactales Callitrichales Calycanthales Calycerales Campanulales Campynematales Can

ellales Cannales CapparalesCaprifoliales Cardiopteridales Carduales Caricales Carle

manniales Caryophyllales Cassiales Casuarinales CelastralesCentrolepidales Cephalot

ales Cephalotaxales Ceratophyllales Cercidiphyllales Chenopodiales Chironiales Chlor

anthalesChrysobalanales Cinchonales Circaeasterales Cistales Citrales Cocosales Col

chicales Columelliales CombretalesCommelinales Connarales Convolvulales Coriarial

es Cornales Corylales Corynocarpales Crassulales CrossosomatalesCucurbitales Cunn

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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inghamiales Cunoniales Cupressales Cycadales Cyclanthales Cymodoceales Cynarales

CynomorialesCyperales Cyrillales Cytinales

Daphnales Daphniphyllales Dasypogonales Datiscales Degeneriales Desfontainiales D

iapensiales Didymelales DillenialesDioales Dioncophyllales Dioscoreales Diospyrales

Dipentodontales Dipsacales Droserales

Ebenales Echiales Elaeagnales Elaeocarpales Elatinales Elodeales Emmotales Empetr

ales Ephedrales Ericales EriocaulalesErythropalaless Escalloniales Eucommiales Euph

orbiales Eupomatiales Eupteleales Euryalaless

Fabales Fagales Falcatifoliales Ficales Flacourtiales Flagellariales Fouquieriales Franc

oales Frangulales

Galiales Garryales Geissolomatales Gentianales Geraniales Gesneriales Ginkgoales G

laucidiales Globulariales GnetalesGoodeniales Greyiales Griseliniales Grossulariales

Grubbiales Gunnerales Gyrocarpales Gyrostemonales

Haemodorales Haloragales Hamamelidales Hanguanales Heisteriales Helleborales He

lwingiales Himantandrales HippuridalesHomaliales Hortensiales Huales Huerteales

Hydatellales Hydnorales Hydrangeales Hydrastidales HydrocharitalesHydropeltidales

Hydrostachyales Hypericales Hypoxidales

Icacinales Illiciales Iridales Irvingales Iteales Ixiales

Jasminales Juglandales Julianiales Juncaginales Juncales

Lacistematales Lactoridales Lamiales Lardizabalales Laurales Lecythidales Ledocarpal

es Leitneriales LentibularialesLigustrales Liliales Limnanthales Linales Liriales Loasal

es Lobeliales Loganiales Lonicerales Loranthales LowialesLythrales

Magnoliales Malpighiales Malvales Marathrales Marcgraviales Mayacales Medusagy

nales Medusandrales MelanthialesMelastomatales Meliales Melianthales Meliosmal

es Menispermales Menyanthales Metteniusales MitrastemonalesMiyoshiales Moni

miales Moringales Musales Myricales Myristicales Myrothamnales Myrsinales Myrta

les

Najadales Nandinales Narcissales Nartheciales Nelumbonales Nepenthales Neuradal

es Nitrariales Nolanales NothofagalesNyctaginales Nymphaeales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Ochnales Oenotherales Olacales Oleales Onagrales Oncothecales Opuntiales Orchid

ales Oxalidales

Saxifragales Pandales Pandanales Papaverales Paracryphiales Parasitaxales Paridales

Parnassiales PassifloralesPenaeales Pennantiales Pentaphragmatales Peridiscales P

etiveriales Petrosaviales Phellinales Philydrales PhyllanthalesPhysenales Phytolaccal

es Picramniales Pinales Pinguiculales Piperales Pittosporales Plantaginales Platanale

sPlumbaginales Poales Podocarpales Podophyllales Podostemales Polemoniales Pol

ygalales Polygonales PontederialesPortulacales Posidoniales Potamogetonales Primu

lales Proteales

Quercales Quillajales Quintiniales

Rafflesiales Ranunculales Rapateales Resedales Restionales Rhabdodendrales Rhamn

ales Rhinanthales RhizophoralesRhodorales Rhoipteleales Roridulales Rosales Rouss

eales Rubiales Ruppiales Rutales

Sabiales Salicales Salvadorales Samolales Samydales Sanguisorbales Santalales Sapin

dales Sapotales SarracenialesSaxegotheales Saxifragales Scheuchzeriales Sciadopitya

les Scleranthales Scrophulariales Scyphostegiales SedalesSilenales Simmondsiales S

milacales Solanales Sphenocleales Sphenostemonales Stangeriales Stellariales Stemo

nalesStilbales Stylidiales Styracales Surianales

Taccales Tamales Tamaricales Taxales Taxodiales Tecophilaeales Terebinthales Tern

stroemiales Theales TheligonalesThymelaeales Tiliales Tofieldiales Torricelliales Tov

ariales Tribelales Trilliales Trimeniales Triuridales TrochodendralesTropaeolales Turn

erales Typhales

Ulmales Urticales

Vacciniales Vahliales Vallisneriales Velloziales Veratrales Verbenales Viburnales Vinc

ales Violales Viscales VitalesVochysiales

Welwitschiales Winterales

Xanthorrhoeales Ximeniales Xyridales

Zamiales Zingiberales Zosterales Zygophyllales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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ALPHABETICAL LISTING OF ALL FAMILY NAMES OF SEED PLANTS WITH LINKS

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In some cases clicking on a family synonym will send you directly to the subfamily in

which the genus of the basionym is to be found

Abietaceae Abolbodaceae Abrophyllaceae Acalyphaceae Acanthaceae [Acanthochla

mydaceae] Acarnaceaee AceraceaeAchariaceae Achatocarpaceae Achradaceae Achy

ranthaceae Acmopylaceae Acoraceae Acristaceae ActaeaceaeActinidiaceae Actinota

ceae Amentotaxaceae Adenogrammaceae Adoxaceae Aegialitidaceae Aegicerataceae

Aegilopaceae Aeginetiaceaee Aegiphilaceae Aesculaceae Aextoxicaceae Agapanthac

eae Agavaceae AgdestidaceaeAgialidaceae Agrimoniaceae Agrostidaceae Ailanthace

ae Aitoniaceae Aizoaceae Akaniaceae Alangiaceae AlchemillaceaeAldrovandaceae A

lismataceae Alliaceae Allioniaceae Allophylaceae Aloaceae Alopecuraceae Alpiniacea

e AlseuosmiaceaeAlsinaceae Alsodeiaceae Alstroemeriaceae Altingiaceae Alzateacea

e Amaranthaceae Amaryllidaceae Amborellaceae Ambrosiaceae Amentotaxaceae A

mmanniaceae Ammiaceae Amomaceae Ampelidopsaceae Amygdalaceae Amyridacea

eAnacardiaceae Anagallidaceae Anarthriaceae Anchusaceae Ancistrocladaceae Andr

omedaceae AndropogonaceaeAndrostachyaceae Androsynaceae Anemarrhenaceae

Anemonaceae Angelicaceae Anisophylleaceae AnnonaceaeAnomochloaceae Anopter

aceae Anrederaceae Anthemidaceae Anthericaceae Anthobolaceae Antidesmataceae

Antirrhinaceae Antoniaceae Aparinaceae Aphanopetalaceae Aphloiaceae Aphyllanth

aceae Apiaceae ApocynaceaeApodanthaceae Aponogetonaceae Aporusaceae Apose

ridaceae Apostasiaceae Aptandraceae AquifoliaceaeAquilariaceae Aquilegiaceae A

raceae Aragoaceae Araliaceae Aralidiaceaee Araucariaceae Arbutaceae Arceuthidace

aeArceuthobiaceae Arctostaphylaceae Arctotidaceae Ardisiaceae Arecaceae Argoph

yllaceae Arisaraceae AristolochiaceaeAristoteliaceae Arjonaceae Armeriaceae Artem

isiaceae Arthrotaxidaceae Artocarpaceae Arundinaceae ArundinellaceaeAsaraceae A

sclepiadaceae Ascyraceaee Asiraceae Asparagaceae Asperulaceae Asphodelaceae As

pidistraceaeeAsteliaceae Asteraceae Asteranthaceae Asteropeiaceae Astragalaceae

Astrocarpaceae AthanasiaceaeAtherospermataceae Atriplicaceae Atropaceae Aucuba

ceae Aurantiaceae Austrobaileyaceae Austrotaxaceae AvenaceaeAverrhoaceae Avet

raceae Avicenniaceae Azaleaceae Azimaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Balanitaceae Balanopaceae Balanophoraceae Balsameaceae Balsaminaceae Bambusa

ceae Barbaceniaceae BarbeuiaceaeBarbeyaceae Barclayaceae Barringtoniaceae Base

llaceae Bataceae Baueraceae Bauhiniaceae Baxteriaceae BegoniaceaeBehniaceae B

elangeraceae Belloniaceae Bembiciaceae Berberidaceae Berberidopsidaceae Berryac

eaee BersamaceaeBertyaceae Berzeliaceae Besleriaceae Betaceae Betulaceae Biebe

rsteiniaceae Bifariaceae Bignoniaceae BischofiaceaeBixaceae Blakeaceae Blandfordia

ceae Blattiaceae Blepharocaryaceae Blitaceae Blyxaceae Boerlagellaceae Bolivaracea

eBombacaceae Bonnetiaceae Bontiaceae Boopidaceae Boraginaceae Borassaceae B

oroniaceae BoryaceaeBotryodendraceae Bougainvilleaceae Boweniaceae Brassicacea

e Brachycaulaceae Bretschneideraceae BrexiaceaeBromeliaceae Brownlowiaceae Br

unelliaceae Bruniaceae Brunoniaceae Brunsvigiaceae Bryoniaceae BuchneraceaeBuci

daceae Buddlejaceae Buglossaceae Bulbocodiaceae Bumeliaceae Bupleuraceae Burc

hardiaceae BurmanniaceaeBurseraceae Butneriaceae Butomaceae Buxaceae Byblida

ceae Byttneriaceae

Cabombaceae Cacaoaceae Cactaceae Caesalpiniaceae Caladiaceae Calamaceae Calce

olariaceae CalectasiaceaeCalendulaceae Callaceae Callicomaceae Calligonaceae Calli

traceae Callitrichaceae Calochortaceae CalophyllaceaeCalthaceae Calycanthaceae Ca

lyceraceae Cambogiaceae Camelliaceae Campanulaceae CampynemataceaeCanacom

yricaceae Canellaceae Cannabaceae Cannaceae Canopodaceae Canotiaceae Cansjera

ceae CapparaceaeCaprariaceae Caprifoliaceae Cardiopteridaceae Carduaceae Carica

ceae Carissaceae Carlemanniaceae CarpinaceaeCarpodetaceae Cartonemataceae Ca

ryocaraceae Caryophyllaceae Casearieaceae Cassiaceae CassipoureaceaeCassythacea

e Castaneaceae Castelaceae Casuarinaceae Catesbaeaceae Cathedraceae Cecropiace

ae CedraceaeCedrelaceae Celastraceae Celosiaceae Celtidaceae Centaureaceae Cen

trolepidaceae Centroplacaceae CepaceaeCephalanthaceae Cephalotaceae Cephalota

xaceae Cerastiaceae Ceratoniaceae Ceratophyllaceae CerberaceaeCercidiphyllaceae

Cercocarpaceae Cercodiaceae Cereaceae Cerinthaceae Ceroxylaceae Cestraceae Cev

alliaceaeChailletiaceae Chamaedoraceae Chamaemoraceae Chamelauciaceae Chaun

ochitonaceae Chelidoniaceae ChelonaceaeChenopodiaceae Chimonanthaceae Chingi

thamnaceae Chionographidaceae Chiranthodendraceae ChironiaceaeChlaenaceae Ch

loanthaceae Chloranthaceae Chloridaceae Chlorogalaceae Chrysobalanaceae Cicerac

eae CichoriaceaeCimicifugaceae Cinchoniaceae Circaeaceae Circaeasteraceae Cissac

eae Cistaceae Citraceae Clematidaceae CleomaceaeClethraceae Clusiaceae Cneorac

eae Cnestidaceae Cnicaceae Cobaeaceae Cochlospermaceae Cocosaceae Coffeaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Colchicaceae Coleogynaceae Columelliaceae Combretaceae Commelinaceae Comocl

adiaceae Compositae CompsoaceaeConantheraceae Connaraceae Conostylidaceae C

onvallariaceae Convolvulaceae Cordiaceae CoreopsidaceaeCoriandraceae Coriariacea

e Coridaceae Corispermaceae Cornaceae Corokiaceae Coronillaceae Correaceae Cor

rigiolaceaeCorsiaceae Corydalaceae Corylaceae Corynocarpaceae Coryphaceae Cost

aceae Cotyledonaceae CoulaceaeCoutariaceae Coutoubiaceae Crassulaceae Crescen

tiaceae Cressaceae Crinaceae Crocaceae CroomiaceaeCrossosomataceae Crotonacea

e Cruciferae Cryptaceae Crypteroniaceae Cryptocorynaceae CryptomeriaceaeCtenol

ophonaceae Cucurbitaceae Cunninghamiaceae Cunoniaceae Cupressaceae Curcumac

eae Curtisiaceae CuscutaceaeCyananthaceae Cyanastraceae Cynanchaceae Cyanella

ceae Cycadaceae Cyclanthaceae CyclantheraceaeCyclocheilaceae Cydoniaceae Cymo

doceaceae Cynaraceae Cynocrambaceae Cynomoriaceae Cyperaceae CyphiaceaeCyp

hocarpaceae Cypripediaceae Cyrillaceae Cyrtandraceae Cyrtanthaceae Cytinaceae

Dacrycarpaceae Dactylanthaceae Damasoniaceae Daphnaceae Daphniphyllaceae Das

ypogonaceae DatiscaceaeDaturaceae Daucaceae Davidiaceae Davidsoniaceae Decais

neaceae Deeringiaceae Degeneriaceae DelphiniaceaeDendrophthoaceae Columelliac

eae Detariaceae Dialypetalanthaceae Dianellaceae Dianthaceae DiapensiaceaeDicha

petalaceae Dichondraceae Diclidantheraceae Dicrastylidiaceae Dictamnaceae Didiere

aceae DidymelaceaeDidymocarpaceae Diegodendraceae Diervillaceae Digitalidaceae

e Dilatridaceae Dilleniaceae Dioaceae DionaeaceaeDioncophyllaceae Dioscoreaceae

Diosmaceae Diospyraceae Dipentodontaceae Diphylleiaceae Diplolaenaceae Dipsaca

ceaeDipterocarpaceae Dirachmaceae Disanthaceae Diselmaceae Dodonaeaceae Do

mbeyaceae Donatiaceae DorsteniaceaeDortmannaceae Doryanthaceae Drabaceae D

racaenaceae Dracontiaceae Drimyidaceae Droseraceae DrosophyllaceaeDryadaceae

Duabangaceae Duckeodendraceae Dulongiaceae Durantaceae Durionaceae Dysphani

aceae

Ebenaceae Eccremidaceae Ecdeiocoleaceae Echiaceae Echinopaceae Ehretiaceae Ela

eagnaceae ElaeocarpaceaeElegiaceae Elatinaceae Ellisiaceae Ellisiophyllaceae Elodea

ceae Elytranthaceae Embeliaceae Emblingiaceae EmmotaceaeEmpetraceae Encepha

lartaceae Engelhardtiaceae Enhalaceae Epacridaceae Ephedraceae Ephemeranthacea

eEpilobiaceae Epimediaceae Eragrostidaceae Eremolepidaceae Eremosynaceae Erica

ceae Erinaceae EriocaulaceaeEriogonaceae Eriospermaceae Erodiaceae Erycibaceae

Eryngiaceae Erysimaceae Erythroniaceae ErythropalaceaeErythrospermaceae Erythro

xylaceae Escalloniaceae Eschscholziaceae Eucomidaceae Eucommiaceae Eucryphiace

aeEuonymaceae Eupatoriaceae Euphorbiaceae Euphrasiaceae Euphroniaceae Eupo

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matiaceae Eupteleaceae EuryalaceaeEustrephaceae Euthemidaceae Eutocaceae Exb

ucklandiaceae Exocarpaceae

Fabaceae Fagaceae Falcatifoliaceae Ferulaceae Festucaceae Ficaceae Ficoidaceae Fit

zroyaceae FlacourtiaceaeFlagellariaceae Flindersiaceae Foetidiaceae Forestieraceae

Fothergillaceae Fouquieriaceae Fragariaceae FrancoaceaeFrangulaceae Frankeniacea

e Fraxinaceae Fraxinellaceae Freycinetiaceae Fritillariaceae Fuchsiaceae Fumariacea

eFunkiaceae

Gaiadendraceae Galacaceae Galanthaceae Galaxiaceae Galedupaceae Galeniaceae G

aliaceae Garciniaceae GardeniaceaeGardneriaceae Garryaceae Geissolomataceae Ge

itonoplesiaceae Gelsemiaceae Geniostomaceae GentianaceaeGeonomataceae Geosir

idaceae Geraniaceae Gerrardinaceae Gesneriaceae Gethyllidaceae Gilliesiaceae Gina

lloaceaeGinkgoaceae Gisekiaceae Gladiolaceae Glaucidiaceae Glechomaceae Glinace

ae Globulariaceae Gnaphaliaceae GnetaceaeGoetzeaceae Gomortegaceae Gomphiac

eae Gomphrenaceae Gonystylaceae Goodeniaceae Gordoniaceae GouaniaceaeGoup

iaceae Gramineae Gratiolaceae Grewiaceae Greyiaceae Grielaceae Griseliniaceae Gr

onoviaceae GrossulariaceaeGrubbiaceae Guaiacanaceae Guamatelaceae Guettardac

eae Gunneraceae Gustaviaceae Guttiferae GyrocarpaceaeGyrostemonaceae

Hachetteaceae Haemanthaceae Haemodoraceae Halesiaceae Hallieraceae Halocarpa

ceae Halophilaceae HalophytaceaeHaloragaceae Hamamelidaceae Hanguanaceae H

aptanthaceae Harmandaceae Hebenstretiaceae HectorellaceaeHederaceae Hedyosm

aceae Hedyotidaceae Hedysaraceae Heisteriaceae Heleniaceae Heliamphoraceae H

elianthaceaeHelianthemaceae Helichrysaceae Heliconiaceae Helicteraceae Heliotropi

aceae Helleboraceae Heloniadaceae HelosidaceaeHelwingiaceae Hemerocallidaceae

Hemimeridaceae Hemiphylacaceae Henriqueziaceae Henslowiaceae Hermanniaceae

Hernandiaceae Herniariaceae Herreriaceae Hesperocallidaceae Heterantheraceae He

teropyxidaceae HeterostylaceaeHibbertiaceae Hibiscaceae Hilleriaceae Himantandra

ceae Hippocastanaceae Hippocrateaceae HippomanaceaeHippophaeaceae Hippurida

ceae Hirtellaceae Holacanthaceae Homaliaceae Hopkinsiaceae Hoplestigmataceae H

ordeaceaeHornschuchiaceae Hortensiaceae Hortoniaceae Hostaceae Houstoniaceae

Huaceae Huerteaceae HugoniaceaeHumbertiaceae Humiriaceae Hyacinthaceae Hyd

atellaceae Hydnoraceae Hydrangeaceae Hydrastidaceae HydrillaceaeHydrocharitacea

e Hydrocotylaceae Hydrogetonaceae Hydroleaceae Hydropeltidaceae Hydrophylacac

eae HydrophyllaceaeHydrostachyaceae Hymenocardiaceae Hypecoaceae Hyperanth

eraceae Hypericaceae Hypopityaceae HypoxidaceaeHypseocharitaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta

ceae ImperatoriaceaeInocarpaceae Inulaceae Ionidiaceae Iriarteaceae Iridaceae Irvi

ngiaceae Isnardiaceae Isophysidaceae IteaceaeStrasburgeriaceae Ixiaceae Ixioliriace

ae Ixonanthaceae

Jalapaceae Jamboliferaceae Japonoliriaceae Jasionaceae Jasminiaceae Johnsoniaceae

Joinvilleaceae JuglandaceaeJulianiaceae Juncaceae Juncaginaceae Juniperaceae Jus

siaeaceae Justiciaceae

Kadsuraceae Kaliphoraceae Kalmiaceae Kaniaceae Kiggelariaceae Kingdoniaceae Kin

giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae

Krameriaceae

Labiatae Lacandoniaceae Lachenaliaceae Lacistemataceae Lactoridaceae Lactucaceae

Lagerstroemiaceaee LamiaceaeLampsanaceae Lanariaceae Langsdorffiaceae Lantan

aceae Lapageriaceae Lardizabalaceae Lasiopetalaceae LauraceaeLawsoniaceae Laxm

anniaceae Lecythidaceae Ledaceae Ledocarpaceae Leeaceae Leguminosae Leitneriac

eae LemnaceaeLennoaceae Lentibulariaceae Lentiscaceae Leoniaceae Leonticaceae

Lepidobotryaceae LepidocarpaceaeLepidocaryaceae Lepidocerataceae Leptaulaceae

Leptospermaceae Lepturaceae Lepuropetalaceae LeuctenbergiaceaeLibocedraceae

Licaniaceae Ligustraceae Lilaeaceae Liliaceae Limeaceae Limnanthaceae Limnocharit

aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace

ae Linderniaceae Lindleyaceae Linnaeaceae LippayaceaeLiriaceae Liriodendraceae Li

ssocarpaceae Littorellaceae Loasaceae Lobeliaceaee Loganiaceae LomandraceaeLoni

ceraceae Lopeziaceae Lophiocarpaceae Lophiolaceae Lophiraceae Lophophytaceae L

ophopyxidaceae LoranthaceaeLotaceae Lowiaceae Lupulaceaee Luxemburgiaceae Lu

zuriagaceae Lychnidaceae Lyciaceae LyginiaceaeLygodisodeaceae Lysimachiaceae Ly

thraceae

Macarisiaceae Mackinlayaceae Madiaceae Maesaceae Magnoliaceae Malaceae Male

sherbiaceae MalortieaceaeMalpighiaceae Malvaceae Mangiaceae Manicariaceae Ma

paniaceae Marantaceae Marathraceae MarcgraviaceaeMartyniaceae Mastixiaceae

Matricariaceae Maundiaceae Mayacaceae Medeolaceae Medusagynaceae Peridiscac

eaeMelaleucaceae Melampyraceae Melanophyllaceae Melanthiaceae Melastomatac

eae Meliaceae Melianthaceae MelicaceaeMeliosmaceae Melittidaceae Melochiacea

e Memecylaceae Mendonciaceae Menispermaceae Menthaceae MenyanthaceaeMe

nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

Nypaceae Nyssaceae

Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

ceae Papilionaceae Pappophoraceae PapyraceaeParacryphiaceae Parasitaxaceae Par

ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

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hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

50

ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 18: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

18

Even although relationships of many parasitic and aquatic groups are being

clarified and at least some of the proposed phylogenetic positions seem to be well

supported it is indeed often still very difficult to understand how such groups have

evolved from their more morphologically conventional relatives However we have had

all sorts of preconceptions as to how characters should evolve Thus we used to

assume that features like highly scalariform vessel perforation plates or the complete

absence of vessels or a flower with an androecium that had many stamens a superior

ovary or separate petals were necessarily plesiomorphic or primitive and conversely

a simple perforation plate and the presence of vessels an androecium with few

stamens inferior ovary or petals that were connnate were almost necessarily

apomorphic or advanced (but cf eg Stebbins 1951) Such assumptions are incorrect

(eg Soltis et al 2005b) Carpels may become secondarily free carpels may fail to close

the seeds then developing outside the confines of the carpel as in some Aspagaraceae-

Nolinoideae Violaceae Berberidaceae Malvaceae-Sterculioideae etc) in Peliosanthes

teta perhaps the only species in Peliosanthes(Asparagaceae-Nolinoideae) the ovary

varies from superior to inferior (Jessop 1976 species limits here need close

investigation see also Kuzoff et al 2001 and Soltis amp Hufford 2002 Saxifragaceae

Apiales etc) many-seeded carpels can evolve from few-seeded carpels

(Razafimandimbison et al 2008) monoecy may be derived from dioecy (Schaefer amp

Renner 2010 and references) Classic studies such as those by Babcock (eg 1947)

on Crepis that assumed that evolution - in this case of the karyotype in particular - was

unidirectional have needed comprehensive re-evaluation (Enke amp Gemeinholzer 2008)

Most if not all characters have reversed andor evolved in parallel as is clear in the

discussion of ovary position here especially in Poales Saxifragales and Asparagales and

also in comments on such morphologically distinctive taxa (when compared with their

immediate relatives) such as Menyanthaceae and especially Pittosporaceae Parallel

evolution may occur even at the level of amino acid substitution as in the independent

acquisition of the phosphoenolpyruvate carboxylase (pepC) gene in C4 photosynthesis in

grasses (Christin et al 2007b see also Blaumlsing et al 2000)

Thinking of how such characters have been used in the past in classifications at the

generic level the results have been similar to those just discussed above for parasitic

plants and others Relying too much on animal pollination syndromes to mark generic

boundaries has all too often led to taxa that are highly unsatisfactory phylogenetically

(see eg Acanthaceae Bignoniaceae Campanulaceae Ericaceae Lamiaceae

Orchidaceae) and over-reliance on characters of fruit and seed (see eg Brassicaceae

and Apiaceae) has also led to unsatisfactory generic limits Again the more general

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

19

problem is the use of one or a very few characters that have been weighted a priori to

structure classifications (see also Garciacutea et al 2009) although it is perhaps not

surprising that such an approach sometimes results in the delineation of taxa that have

indeed turned out to be quite distinct (cf eg Burtt Astragalus versus Oxytropis)

Taxa recognized here are monophyletic yet many are polythetic at the

morphological level that is they lack unique features characterizing (defining) all and

only members of that family they can be recognised phenetically only by the unique

combinations of characters that they posess This is the result of evolution any

synapomorphy characterizing a taxon may be lost or modified beyond easy recognition

in some of its members or the synapomorphy may appear to be identical to a feature

that has evolved in parallel in a quite unrelated plant That plant groups are polythetic is

almost as much a feature of monophyletic taxa as the paraphyletic taxa common in

evolutionary classifications Some families now include substantial variation as

phenetically distinct derived groups are placed in their proper phylogenetic position -

examples are Ericaceae which include the erstwhile Empetraceae a wind-pollinated

group that is florally very different and the overwhelmingly large-flowered and animal-

pollinated Plantaginaceae which now include several derived small-flowered aquatic

and wind-pollinated groups that were previously placed in separate families

Thus some families as delimited here may not be easy to recognize However

remember that detecting relationships - use whatever characters you can even if they

are not obvious - and naming a plant - focus on easy-to-see characters that may not

reflect relationships - are quite different problems Taxa although natural may not be

readily recognizable indeed it was in exactly this context that Lamarck worked out the

basic principles of writing dichotomous keys in 1778 (see Scharf 2007 for keys and the

like) Of course Lamarcks idea of nature was very different from ours - he thought that

there was some kind of continuum of form on which living organisms were to be

situated with no real gaps anywhere - but this meant that his genera (for example)

might well not be sharply distinct from each other even if each were part of the real

continuum that was life Lamarcks separation of the establishment of relationships and

groupings in a system and identification of the plants in that system have been widely

accepted (but cf eg Godfray amp Knapp 2004 many of the contributors to Heywood et

al 2007) Perhaps the best way of identifying plants at the family level is by well-made

multiple access keys as in Watson and Dallwitz (1992a onwards family limits there may

differ substantially from those adopted here) Multiple access keys free users from the

constraints of dichotomous keys in which particular characters are needed at each step

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

20

of the identification process before there can be further progress Instead those

characters that happen to be evident on a specimen can be used in whatever order is

convenient when linked to illustrations glossaries etc their power is enormous (see

Dallwitz et al 2000 [2006] for the principles underlying their construction and use)

Nevertheless dichotomous keys such as those of Hutchinson (1973) and Franz Thonner

(Geesink et al 1981) have their uses Of course taxa coming out adjacent in keys may

well not be at all related

(When identifying large numbers of plants even more efficient than either style of

identification and certainly lots more fun is sight identification Unless you have a

photographic memory you have to build up your knowledge of comparative plant

morphology - on which the ability to make accurate identifications depends - by

repeated observation When faced with an unknown plant I always look for leaf teeth

and stipules distinctive hair types especially stellate and T-shaped hairs glands and

punctations of any sort the presence of latex or other exudate and check leaf insertion

smelling crushed leaves can also be helpful In this context nodal anatomy can usually

be checked using a razor and a hand lens or even simply carefully examining leaf scars

The short paragraphs added after most families may help in confirming familial

identifications)

For the record and for the little that it is worth there are 4 orders and 13 families

of gymnosperms characterised on these pages and together they include some 82

genera and 947 species For angiosperms comparable figures are 56 orders 445

families 13208 genera and 261750 species (of which monocots include 11 orders 89

families 2759 genera and 52760 species) Note however that higher mathematics

was never my strong point and anyway these are pretty meaningless figures even for

species which many (but not all) might concede smacked slightly more of reality than

other taxa estimates range as high as 422000 (Govaerts 2001) Furthermore numbers

of genera and species change daily Nevertheless as emphasized here families are

useful in teaching we as a community can ensure that their limits remain largely stable

and by concentrating on relatively few of them one can gain some familiarity with much

of the worlds flora For summaries of the sizes of orders and families and of the general

arrangement followed in these pages see also the Statistics page itself

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

21

A PHYLOGENETIC TREE OF SEED PLANTS (THE MAIN TREE)

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

22

The Main Tree above is a conservative summary of well supported relationships

between the major clades of extant seed plants Reading - interpreting and

understanding - such a phylogenetic tree is not necessarily easy so reading a tree may

require a bit of practice OHara (eg 1988 1992) provides an accessible introduction to

thinking and talking about history in the context of looking at phylogenetic trees More

recently Krell and Cranston (2004) Crisp and Cook (2005) and others have emphasized

how careful one must be when interpreting and talking about evolution in ladderized

trees in particular and phylogenetic trees in general Gregory (2008) provides a

particularly detailed discussion of this problem In ladderized trees the smaller (in terms

of numbers of terminals) sister taxon is consistently shown on the same side at every

node thus the trees tend to be pectinate like the teeth of a comb however the

horizontal axis that is as a result so evident has no polarity or particular direction The

use of the adjective basal when discussing phylogenetic trees is especially dangerous

(see also Wojciechowski et al 2004 D Soltis et al 2005b) When I use the term and the

context is not otherwise clear I am referring to the pectinations at the base of a

ladderised tree Remember that when talking about sister taxa one can never be basal

to the other or older or younger although members of one can be more derived - in the

sense of having more apomorphies - than the other (but even then one has to be

careful) The words primitive and advanced and lower and higher should

practically never be used especially when talking about taxa Amborellaceae and

Pinaceae are sister to all other flowering plants and Pinales and are basal to the crown

groups of all other flowering plantsPinales respectively but that does not mean they

are thereby primitive Pinaceae in particular have numerous apomorphies The word

plesiomorphic is far less loaded than primitive and can be used to talk about

individual characters

Most trees here have been more or less ladderized as is the one above The

ladderization may be imperfect for example you can see that on the Main Tree

asterids with ten orders follow rosids and relatives which have seventeen although

this is in part because of the currently unresolved nature of relationships between the

main clades of core eudicots (this is likely to change soon) The Main Tree could have

been drawn with Amborella Acorus or a host of other taxa at the far right without

offending any relationships phylogenetic trees are like mobiles the only fixed points

being the nodes Despite the problems just mentioned pectination interpreted

carefully has its value and it is not an arbitrary process As one reads the terminals of a

pectinate tree from left to right adjacent terminals will be separated by apomorphies

that will cumulate in an appropriate fashion When thinking of a book or a herbarium

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

23

sequence (see Haston et al 2007) this is of some value Nymphaeales and

Austrobaileyales are here adjacent on the tree but they could be separated by

hundreds of families in the sequence merely by rotating only the node from which

Nymphaeales and their sister group (all other angiosperms minus about 75 species)

arise If adjacent in a book or herbarium or as in these summary pages then it is

relatively easy to relate their characters both apomorphic and plesiomorphic but if

separated by hundreds of pages or two floors in a large building then it is less easy to

get anything from the sequence (Since all orders in the pages of the main part of this

site are preceded by the apomorphies of all nodes immediately below them in the seed

plant phylogeny and because of the linkages that have been built in to the site this

problem is much less serious there there is no sequence) So the particular sequence of

all seed plants that is based on a phylogenetic classification but is to be used in herbaria

or any other place where taxa have to be arranged linearly as in this part of APweb

will be one that maximizes the number of taxa that are both successive branches of the

tree and placed successively in the sequence Since specimens are generally filed under

families the outline of a new family sequence for arranging herbaria and books can now

be suggested (Haston et al 2007) and such a standardized sequence will further help

teaching and learning about plants

ON THINKING ABOUT APOMORPHIES

Identifying apomorphies is important because understanding the evolution of

morphology in the broad sense - ie including anatomy chemistry etc - in the context

of diversification is one of our major goals For identifying apomorphies several

preconditions must be met One needs to have an accurate well supported phylogeny

one has to have examined the right taxa both from the point of view of morphology and

molecules one has to have coded the characters right (ie delimited states

appropriately) and one has to use an appropriate model of evolution when fitting the

variation to the tree (see eg Omland 1999 Stevens 2006b) If the distributions of

apomorphies below are compared with those in other studies (eg Turgeon et al 2001

Bremer et al 2001 Endress 2001 Albach et al 2001a Judd amp Olmstead 2004 D Soltis

et al 2005b Zhang et al 2006 Judd et al 2007) the reader may find a number of

differences Although I have integrated such studies as far as is possible there are five

reasons that link to the preconditions just mentioned why there may be differences

where features are placed on trees

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

24

1 I may not have found all the information about a particular character there may be

disagreement over its interpretation or I have added information

2 The sampling of nearly all molecular studies is very incomplete (see Salisbury amp Kim

2001 for problems caused by sampling) indeed it is commonplace to decry the

incompleteness of molecular sampling But not only is the sampling in molecular studies

often less than we might wish that of the morphological and chemical characters whose

evolution we are interested in understanding is also often very poor So for many

anatomical chemical and embryological characters that are confidently said to

characterise families and other groups we all too often have no idea if those characters

are applicable to the whole clade or just to a subgroup within that clade However as

our sampling improves we can locate changes on the tree more precisely if in

unexpected positions Thus is is sometimes suggested that Ericaceae have ellagic acid

(Soltis et al 2005b) in fact the little evidence we have indicates that only a small clade

of some 80 species has ellagic acid and the rest of the family - itself alone about one

third of all Ericales - does not Similarly Ericaceae are noted for their

ectendomycorrhizal associations but Enkianthus sister to all other Ericaceae

apparently lacks these (Abe 2005) Thus these mycorrhizae and their associated

distinctive hair roots are unlikely to be apomorphies of Ericaceae - however it appears

that other families near Ericaceae may also have ectendomycorrhizae (see Asai 1934)

3 I am fitting characters to a very conservative tree with many polytomies although the

nodes that are utilised are for the most part strongly supported polytomies make the

optimisation of characters that is the assigment of character state change to a

particular node on the tree notably difficult (eg Madison amp Madison 2002) In nearly

all studies of the evolution of characters distributions of characters are optimised on a

more or less fully resolved tree and the construction of supertrees may yield yet more

detailed hypotheses of relationships (for literature on supertrees see Cotton amp

Wilkinson 2007 2008) Of course some nodes on such fully resolved trees andor

supertrees may have little support and optimisations of characters on such trees may

carry correspondingly little conviction Even parts of some of the trees used here have

poor support eg relationships within aquatic Alismatales etc although I indicate

when this is the case

4 Exactly how one goes about optimising a character on a tree is critically important

Even using simple parsimony optimisations (ACCTRAN or DELTRAN ACCelerated

TRANsitions or DELayed TRANsitions) the position of synapomorphies on trees - and

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

25

hence our ideas of evolution - may differ and this is still more true if one uses maximum

likelihood or Bayesian methods and makes apparently reasonable suggestions about

weighting gains over losses (eg Donoghue amp Ackerley 1996 Cunningham et al 1998

Omland 1997 1999 Ree amp Donoghue 1999 Polly 2001 Webster amp Purvis 2001

Ronquist 2004 Crisp amp Cook 2005) Sannier et al (2007) show how in in Arecaceae that

where on a phylogenetic tree one might peg changes in microsporogenesis will depend

on the methods one uses to do this and Pedersen et al (2007) discuss the sometimes

very substantial effect of node support on the posterior probabilities of ancestral

character states Here I use parsimony optimization not always as explicit as it might be

but I have often indicated where there are particularly important uncertainties as to the

positions of particular character changes on the tree

5 Finally although I have paid quite a lot of attention to the delimitation of the

character states that make up all the characterizations I have not spent enough time on

this critical operation If we are interested in understanding evolution then fitting the

basic variation - not character states - to a tree in principle allows greater flexibility in

understanding morphology in the context of local phylogenies (see also Stevens 2000

Endress 2005c) However many character states used here are delimited globally that

is they are circumscribed in the context of the variation shown by individual characters

across all angiosperms andor in the context of classic ideas of character evolution

Character states often have arbitrary limits and serve best to communicate

information whether they are in fact suitable for either phylogenetic analysis or

understanding evolution are separate issues Studies have rather unsurprisingly

perhaps but importantly shown that dividing the one character into different sets of

states may yield differing ideas of evolution of that character (eg Lamb Frye amp Kron

2003 Hibbett 2004) When looking at trees on which character states are optimised

one should bear in mind the problems surrounding the delimitation of states (eg

Stevens 2000 2006b) and the danger of using pollen or other types - constructs

based on many characters that vary independently but which effectively get lost in

these types thus Blackmore et al (2009) decomposed the pollen types in Asteraceae

into 52 characters

I use Remanes three main criteria of homology or better similarity when

determining the basic similarity of structures on different organisms (see Remane 1952)

These criteria are special properties position and intermediates Special properties

include anatomical or chemical characters gene expression data etc Position refers

to the position of an organ with respect to landmarks on the plant Although plants are

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

26

plants and landmarks are not as fixed as one might like trying to work out the relative

positions of parts is a good way of understanding morphology so long as one does not

try to out-do Procrustes Intermediates include intermediates found on the same

plant or on different plants Intermediates may observed during development or be

adult structures Thus one may find changes during development which suggest what

the nature of a particular structure is and very different stuctures on different plants

may turn out to be similar early in development Or one can simply compare different

taxa and see that structures that appear to be very different when comparing two

different taxa can be linked morphologically by looking at other taxa

Nevertheless the use of these criteria may not yield an unambiguous answer as to

what a structure is even given a solid phylogeny and an improved understanding of

development (see Jaramillo amp Kramer 2007 for a useful discussion) As Endress (2005c)

observed a number of features - position function development shape anatomy

histology gene activity and relationships to other taxa that clearly have petals - can be

used to distinguish a petal (for example) from other floral structures if a petal does not

have one of these features is it thereby not a petal Thus Maturen et al (2005) recently

found that floral organ diversity genes (B and C) were expressed in the large white

inflorescence bracts of Cornus (see also Costa et al 2005) Peney et al (2005) noted that

not all monosulcate pollen grains in monocots have the same developmental pathway

and that as a result such pollen might not have the same ancestral state Reeves and

Olmstead (2003) suggested that the genetic mechanisms causing monosymmetry in

Lamiales and Solanales were different and Serna and Martin (2006) described similar

problems with the development of hairs in Arabidopsis when compared with that of

hairs in Antirrhinum and Solanaceae Indeed as one perhaps might expect delimitation

of states and characters does not necessarily become easier with increasing knowledge

of development etc Thus Buzgo et al (2004) Matthews and Endress (2005) and others

have shown how hard it can be to distinguish between eg prophylls and other floral

structures as their behaviour is studied during the course of the development of the

flower Similarly at what concentration is a particular secondary metabolite deemed to

be present (Waterman (2007)

The validity of the approach used here that of fitting morphological variation to a

largely molecular-based tree may be questioned However I think it rather unlikely that

well-supported molecular branches will be overturned by morphological data Indeed

analyses of morphological data alone do provide support for many of the clades evident

in molecule-only analyses and analyses with morphological and molecular data

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

27

together may lead to increased support for clades (eg Hufford 1992 Nandi et al 1998

[but adding morphological data reduces support for a number of critical clades too]

Doyle amp Endress 2000 however in none of these papers is the use of morphology

without ambiguity) It is unfortunately clear that the use of morphology alone may not

suggest problems in the phylogenetic placement of taxa that later turn out to have been

wrongly included (eg Zhang et al 1992) Thus I have been wary of putting much weight

on clades that have only morphological support but note that I have treated molecule-

based clades with low bootstrap or jacknife support values (esp below 70) or low

posterior probabilies (below 095) likewise Although I may have been mistaken in

placing so much emphasis on molecular data in terms of providing the basic

phylogenetic framework for angiosperms morphological and molecular data are only

very rarely in irreconcilably strong conflict There are indeed a few places where the

conflict seems extreme These include the relative positions of the Monimiaceae and

Hernandiaceae (Laurales) the position of Hanguanaceae (Commelinales [as here] or

Zingiberales) and of Triplostegia (is it in Dipsacaceae or Valerianaceae - see

Dipsacales) Fossils are unlikely to affect the topologies of the trees presented here but

see below for their importance in understanding morphological evolution in general and

the evolution of angiosperms in particular Nevertheless some largely reject the idea

that trees based on molecular data alone can recover phylogenetic relationships

especially when branching points are old and prefer to used trees based on analysis of

morphological data including those taken from fossils (Hilton amp Bateman 2006 Farjon

2007)

All in all however the extent of the congruence between morphological and

molecular data is impressive and heartening and many clades can be characterised

morphologically It seemed in 1998 that there were no unambiguous morphological

synapomorphies for angiosperm orders (K Bremer 2000) and this is still true if by

unambiguous is meant non-homoplasious However many orders and other clades

have synapomorphies even if these may be indistinguishable at least at the current

level of morphological and developmental knowledge from parallel occurences

elsewhere As our morphological knowledge increases so too does the number of

apomorphies

Endress and Matthews (2006a) emphasize the importance of tendencies and

developmental constraints when thinking about characters of clades within the rosids -

many characters or character combinations occur in a rather sporadic fashion within

the clade and apparently notably less frequently outside it There are a number of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

28

examples of tendencies here and in the individual order pages such as the features

enclosed by parentheses in groups above the level of family eg see (cuticular waxes

as aggregated rodlets) for the commelinids and the discussion of the distribution of

polyandry that is flowers with numerous stamens in the asterid I + II groups etc

Tendencies may involve more than single characters When thinking of the

associations of features like integument thickness and vascularization of the integument

(Endress amp Matthews 2006a) one can imagine fairly simple developmental

preconditions being operative As Endress and Matthews (2006a) note it is difficult to

imagine vascular tissue developing in an integument only two cells thick Changes here

would be loosely correlated if morphologically linked However the strongly correlated

changes noted by Givnish et al (2005) are ecologically linked but are presumably

morphologicallydevelopmentally independent When there seem to be characters

evolving more or less together tests can be carried out to see if the changes are

concentrated on certain branches of the tree (eg Maddison 1990 Sanderson 1991

Maddison amp Maddison 2000) Such tests have rarely been carried out at higher levels in

angiosperms

One of the most striking examples of a tendency is the distribution of N-fixation

restricted as it is to a monophyletic group of four clades although it has arisen seven (or

perhaps several more) times independently within the N-fixing clade and several

members of two quite different kinds of bacteria are involved (eg Clawson et al 2004

Elliott et al 2007 Sprent amp James 2007) Other examples of work that bears on the issue

of tendencies include the findings that flowers of polysymmetrical Arabidopsis have

genes like TCP1 that are expressed asymmetrically during early development and TCP1

is a probable orthologue of the well-known CYC gene of Antirrhinum that is involved in

the development of monosymmetric slowers there (Cubas et al 2001 Costa et al 2005

etc) Parallelism might build on this underlying morphologically cryptic monosymmetry

even if details of the genetic mechanisms causing the monosymmetry evident in

particular groups may be different (Reeves amp Olmstead 2003 Cubas 2004 see above)

Similarly there have been several recent suggestions that the capability to synthesise a

particular metabolite may be switched off but not lost and so can sometimes be

reacquired (eg Wink amp Witte 1983 Wink 2003 Liscombe et al 2005 Larsson 2007

Waterman 2007) Hence perhaps the rather spotty distribution of many secondary

metabolites like ellagic acid the indole alkaloid camptothecin iridoids etc when

considered in the context of phylogenies As a non-botanical example - but a rather nice

one - Salwini-Plawen and Mayr (1961) suggested some time ago that there has been

considerable parallelism (40-65 or more independent origins) in the evolution of eyes in

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

29

metazoans However Pax 6 seems to be a master control gene that is involved in eye

formation perhaps throughout all bilateralians (eg Gehring amp Ikeo 1999 Erwin amp

Davidson 2002) which changes the nature of the problem Note however that in

general our invocation of evolutionary tendencies to explain the patterns of

distributions of characters is really an admission that we do not understand those

patterns

As we find out more about variation we will probably find fewer and fewer features

constant throughout a group Most unqualified statements of presence and absence

should properly be qualified as usually present or usually absent if one is thinking of

the characterisations on the order pages as encompassing the total variation within a

clade Thus Pistia alone among monocots as so far known has sieve tube plastids with

starch grains not protein crystals However this does not affect the fact that sieve tube

plastids with cuneate protein crystals and lacking starch grains are an apomorphy for

monocots Furthermore fossils in a number of cases suggest character combinations

unknown in extant taxa as may be seen in the discussions of Fagaceae Platanaceae

Iteaceae Calycanthaceae etc Confusing the issue there may be questions as to where

exactly on the tree a particular fossil is to be placed (eg see Nymphaeaceae

Calycanthaceae Archaefructus etc)

To summarize given our current understandings of both phylogenies and

characters evolution of some characters in which we are interested seems very labile

(see eg D Soltis et al 2005b Endress and Matthews 2006a Stevens 2006b Ekman et

al 2008 for a good recent study) and I have been cautious when talking about character

evolution Much effort must continue to be spent in summarizing characters of well-

established clades at all levels providing features by which they may be recognized and

signaling synapomorphies Remember that (1) the basic morphological anatomical and

chemical knowledge of many critical taxa is woefully incomplete (2) different

assumptions about character evolution may greatly affect the position of

synapomorphies on trees (3) in many cases relationships within and between many

groups are too uncertain at present to worry very much about synapomorphies and

(4) we must be clear about what we do and do not not know As mentioned above I

have indicated in a number of places where there are particularly important

uncertainties as to where characters should be placed on the tree Nevertheless it is a

relatively easy matter to update notes such as these and it can be a simple matter to

incorporate new data on characters that have never before been considered in the

context of a tree Much basic - and unfortunately perhaps unfashionable - work must be

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

30

carried to clarify the distribution of morphological anatomical and chemical characters

for examples of what can to be done see the work of P K Endress and collaborators

(floral morphology and development) and S R Jensen and collaborators (iridoids) But

acquiring information about nodal anatomy cell and tissue distribution nodal anatomy

and the like is not difficult at all and should be generally encouraged

But all this is merely a necesary prelude to the understanding of evolution For this

one needs to know a lot more including dating the phylogeny understanding the

function(s) of characters factoring in relevant aspects of the palaeoenvironment in

which an apomorphy first appeared etc

SUMMARY OF THE SYSTEM

Below is a formalised summary of the relationships within orders of the families of seed

plants There are a few families that are not recognised even as options in APG II and

vice versa as well as a few extra orders The families to be recognised in the most

recent edition of Mabberleys The Plant Book (Mabberley 2008) are also largely

consistent with those below But all differences are trivial and will - I hope - eventually

disappear

Square brackets - [] - enclose clades the plus sign - + - designates sister taxa a

comma - - denotes part of a polytomy and quotation marks - - denotes a

paraphyletic group

SEED PLANTS

GYMNOSPERMS

Cycadales

Cycadaceae + Zamiaceae

Ginkgoales

Ginkgoaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

31

Pinales

Pinaceae [[Araucariaceae [Podocarpaceae + Phyllocladaceae]] [Sciadopityaceae

[Taxaceae + Cupressaceae]]]

Gnetales

Ephedraceae [Gnetaceae + Welwitschiaceae]

ANGIOSPERMSFLOWERING PLANTS

Amborellales

Amborellaceae

Nymphaeales

Hydatellaceae [Cabombaceae + Nymphaeaceae]

Austrobaileyales

Austrobaileyaceae [Schisandraceae + Trimeniaceae]

Chloranthales

Chloranthaceae

MAGNOLIIDS

Magnoliales

Myristicaceae [Magnoliaceae [[Himantandraceae + Degeneriaceae] [Eupomatiaceae +

Annonaceae]]]

Laurales

Calycanthaceae [[Siparunaceae [Gomortegaceae + Atherospermataceae]] [Monimiaceae

[Hernandiaceae + Lauraceae]]]

Canellales

Canellaceae + Winteraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

32

Piperales

[Hydnoraceae + Aristolochiaceae] [Piperaceae + Saururaceae]

MONOCOTS

Acorales

Acoraceae

Alismatales

Araceae [Tofieldiaceae [[Alismataceae [Hydrocharitaceae + Butomaceae]]

[Scheuchzeriaceae [Aponogetonaceae [Juncaginaceae [Maundiaceae [[Posidoniaceae

[Ruppiaceae + Cymodoceaceae]] [Zosteraceae + Potamogetonaceae]]]]]]]

Petrosaviales

Petrosaviaceae

Dioscoreales

Nartheciaceae [[Taccaceae + Thismiaceae] [Burmanniaceae + Dioscoreaceae]]

Pandanales

Velloziaceae Triuridaceae Stemonaceae [Pandanaceae + Cyclanthaceae]

Liliales

Corsiaceae [Campynemataceae [Petermanniaceae [Colchicaceae + Alstroemeriaceae]]

Melanthiaceae [[Philesiaceae + Rhipogonaceae] [Smilacaceae + Liliaceae]]]

Asparagales

Orchidaceae [[Boryaceae [Blandfordiaceae [Lanariaceae [Asteliaceae + Hypoxidaceae]]]]

[[Ixioliriaceae + Tecophilaeaceae] [Doryanthaceae [Iridaceae [Xeronemataceae

[Xanthorrhoeaceae [Alliaceae + Asparagaceae]]]]]]]

COMMELINIDS

Unplaced

Dasypogonaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

33

Arecales

Arecaceae

Poales

[Typhaceae + Bromeliaceae] [Rapateaceae [[Thurniaceae [Juncaceae + Cyperaceae]]

[[Mayacaceae [Eriocaulaceae + Xyridaceae]] [Flagellariaceae [Anarthriaceae

[Centrolepidaceae + Restionaceae]]] [Joinvilleaceae [Ecdeiocoleaceae + Poaceae]]]]]]]

Commelinales

[Commelinaceae + Hanguanaceae] [Philydraceae [Haemodoraceae + Pontederiaceae]]

Zingiberales

Musaceae [Strelitziaceae + Lowiaceae] Heliconiaceae [[Cannaceae + Marantaceae]

[Costaceae + Zingiberaceae]]

Ceratophyllales

Ceratophyllaceae

EUDICOTS

Ranunculales

Eupteleaceae [Papaveraceae [[[Lardizabalaceae + Circaeasteraceae] [Menispermaceae

[Berberidaceae + Ranunculaceae]]]]

Sabiales

Sabiaceae

Proteales

Nelumbonaceae [Platanaceae + Proteaceae]

Trochodendrales

Trochodendraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

34

Buxales

Haptanthaceae [Buxaceae + Didymelaceae]

CORE EUDICOTS

Gunnerales

Gunneraceae + Myrothamnaceae

Dilleniales

Dilleniaceae

Saxifragales

Peridiscaceae [[Paeoniaceae [Altingiaceae [Hamamelidaceae [Cercidiphyllaceae +

Daphniphyllaceae]]]] [[Crassulaceae [Aphanopetalaceae [Tetracarpaeaceae

[Penthoraceae + Haloragaceae]]]] [Iteaceae [Grossulariaceae + Saxifragaceae]]]]

Cynomoriaceae unplaced

Vitales

Vitaceae

ROSIDS

FABIDROSID I

Zygophyllales

Krameriaceae + Zygophyllaceae

Celastrales

Lepidobotryaceae + Celastraceae

Oxalidales

Huaceae [[Connaraceae + Oxalidaceae] [Cunoniaceae [Elaeocarpaceae [Brunelliaceae +

Cephalotaceae]]]]

Malpighiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

35

[Achariaceae [Goupiaceae [Violaceae + Passifloraceae] [Lacistemataceae Salicaceae]]

[Lophopyxidaceae Putranjivaceae] [Ctenolophonaceae [Erythroxylaceae +

Rhizophoraceae]] Linaceae Ixonanthaceae Humiriaceae Irvingiaceae

Centroplacaceae Pandaceae [[Bonnetiaceae + Clusiaceae] [Calophyllaceae

[Hypericaceae + Podostemaceae]]] [Malpighiaceae + Elatinaceae] Ochnaceae

[Peraceae [Rafflesiaceae + Euphorbiaceae]] [Phyllanthaceae + Picrodendraceae]

[Balanopaceae [[Trigoniaceae + Dichapetalaceae] [Chrysobalanaceae +

Euphroniaceae]]] Caryocaraceae

N-FIXING CLADE

Fabales

Quillajaceae [Fabaceae [Polygalaceae + Surianaceae]]

Rosales

Rosaceae [[Barbeyaceae [Elaeagnaceae Dirachmaceae Rhamnaceae]] [Ulmaceae

[Cannabaceae [Moraceae + Urticaceae]]]]

Cucurbitales

Anisophylleaceae [[Corynocarpaceae + Coriariaceae] [Cucurbitaceae [Tetramelaceae

[Datiscaceae + Begoniaceae]]]] Apodanthaceae

Fagales

Nothofagaceae [Fagaceae [[Myricaceae + Juglandaceae] [Casuarinaceae

[Ticodendraceae + Betulaceae]]]]

MALVIDROSID II

pgtGeraniales

Geraniaceae [[Melianthaceae + Francoaceae] [Vivianaceae + Ledocarpaceae]]

Myrtales

Combretaceae [[Onagraceae + Lythraceae] [[Vochysiaceae + Myrtaceae]

[Melastomataceae [Crypteroniaceae [Alzateaceae + Penaeaceae]]]]]

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

36

Crossosomatales

[Staphyleaceae [Guamatelaceae [Crossosomataceae + Stachyuraceae]]] [Aphloiaceae

[Geissolomataceae + Strasburgeriaceae]]

Picramniales

Picramniaceae

Sapindales

Biebersteiniaceae Nitrariaceae [Kirkiaceae [Anacardiaceae + Burseraceae]]

[Sapindaceae [Simaroubaceae Rutaceae Meliaceae]]

Huerteales

Gerradinaceae [Dipentodontaceae + Tapisciaceae]

Malvales

Neuradaceae [Thymelaeaceae [Sphaerosepalaceae Bixaceae [Cistaceae

[Sarcolaenaceae + Dipterocarpaceae]] [Cytinaceae + Muntingiaceae] Malvaceae]]

Brassicales

[Akaniaceae + Tropaeolaceae] [[Moringaceae + Caricaceae] [Setchellanthaceae

[Limnanthaceae [[Koeberliniaceae [Bataceae + Salvadoraceae]] [Emblingiaceae

[Pentadiplandraceae [Gyrostemonaceae Resedaceae Stixis and relatives] Tovariaceae

[Capparaceae [Cleomaceae + Brassicaceae]]]]]]]]

Berberidopsidales

Aextoxicaceae + Berberidopsidaceae

Santalales

Erythropalaceae Olacaceae [[Loranthaceae [Misodendraceae + Schoepfiaceae]]

[Opiliaceae + Santalaceae]] Balanophoraceae unplaced

Caryophyllales

[[Droseraceae [Nepenthaceae [Drosophyllaceae [Ancistrocladaceae +

Dioncophylleaceae]]]]] [[Frankeniaceae + Tamaricaceae] [Polygonaceae +

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

37

Plumbaginaceae]]] [Rhabdodendraceae [Simmondsiaceae [[Asteropeiaceae +

Physenaceae] [[Caryophyllaceae [Achatocarpaceae + Amaranthaceae]]

[Stegnospermataceae [Limeaceae [[Lophocarpaceae [Barbeuiaceae [Aizoaceae

[Sarcobataceae Phytolaccaceae Nyctaginaceae]]]] [Molluginaceae [Halophytaceae

Montiaceae Basellaceae Didiereaceae [Talinaceae [Portulacaceae [Anacampseros etc +

Cactaceae]]]]]]]]]]]]]

ASTERIDS

Cornales

[Cornaceae + Nyssaceae] [Hydrangeaceae + Loasaceae] Hydrostachyaceae [Curtisiaceae

+ Grubbiaceae]

Ericales

[Balsaminaceae [Marcgraviaceae + Tetrameristaceae]] [[Polemoniaceae +

Fouquieraceae] Lecythidaceae [[Sladeniaceae + Pentaphylacaceae] [Sapotaceae

[Ebenaceae [Maesaceae [Theophrastaceae [Myrsinaceae + Primulaceae]]]]]

[Mitrastemonaceae Theaceae [Symplocaceae [Styracaceae + Diapensiaceae]]

[[Sarraceniaceae [Roridulaceae + Actinidiaceae] [Clethraceae [Cyrillaceae +

Ericaceae]]]]]]

LAMIIDASTERID I

Unplaced

Metteniusaceae Oncothecaceae Icacinaceae and some unplaced ex-Icacinaceae

Garryales

Garryaceae + Eucommiaceae

Unplaced

Boraginaceae Vahliaceae

Gentianales

Rubiaceae [Gentianaceae [Loganiaceae [Gelsemiaceae + Apocynaceae]]]

Lamiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

38

Plocospermataceae [Oleaceae [Tetrachondraceae [Calceolariaceae

[Gesneriaceae Peltanthera [Pedaliaceae Carlemanniaceae Martyniaceae Stilbaceae

Plantaginaceae Linderniaceae Bignoniaceae Verbenaceae Lamiaceae Paulowniaceae

Schlegeliaceae Thomandersiaceae Phrymaceae Rehmannia Acanthaceae

Scrophulariaceae Orobanchaceae Byblidaceae Lentibulariaceae]]]]]

Solanales

[Montiniaceae [Sphenocleaceae + Hydroleaceae]] [Convolvulaceae + Solanaceae]

CAMPANULIDASTERID II

Aquifoliales

[[Cardiopteridaceae + Stemonuraceae] [Aquifoliaceae [Helwingiaceae +

Phyllonomaceae]]

Asterales

[Rousseaceae + Campanulaceae] Pentaphragmataceae [[Alseuosmiaceae [Phellinaceae

+ Argophyllaceae]] Stylidiaceae [Menyanthaceae [Goodeniaceae [Calyceraceae +

Asteraceae]]]]

Escalloniales

Escalloniaceae

Bruniales

[Bruniaceae + Columelliaceae]

Apiales

Pennantiaceae [Torricelliaceae [Griseliniaceae [Pittosporaceae [Araliaceae

[Myodocarpaceae + Apiaceae]]]]]

Paracryphiales

Paracryphiaceae

Dipsacales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

39

Adoxaceae [Diervillaceae [Caprifoliaceae [Linnaeaceae [Morinaceae [Dipsacaceae +

Valerianaceae]]]]]

LINKS TO ORDERS AND FAMILIES

[Back to Top]

MAIN GROUPINGS

Asterids commelinids core eudicots asterid 1 asterid

2 eudicots gymnosperms Magnoliophyta monocots N-fixing claderosids seed plants

ALPHABETICAL LISTING OF ALL ORDINAL NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | L | M | N | O | P | Q | R | S | T | U | V | W | X | Z

Abietales Acanthales Acerales Acorales Actinidiales Actinostrobales Adoxales Aescul

ales Agavales Aizoales AkanialesAlismatales Alliales Alseuosmiales Alstroemeriales

Altingiales Amaranthales Amaryllidales Amborellales AmbrosialesAmmiales Amomal

es Ancistrocladales Anisophylleales Annonales Anthobolales Apiales Apocynales Apo

nogetonalesAquifoliales Arales Araliales Aralidiales Araucariales Arecales Aristolochi

ales Athrotaxidales Asarales AsclepiadalesAsparagales Asphodelales Asparagales Ast

eliales Atriplicales Aucubales Austrobaileyales Avenales

Balanitales Balanopales Balanophorales Balsaminales Barbeyales Barclayales Batales

Begoniales BerberidalesBerberidopsidales Betulales Biebersteiniales Bignoniales Bix

ales Boraginales Brassicales Brexiales Bromeliales BrunialesBrunoniales Burmannial

es Burserales Butomales Buxales Byblidales

Cactales Callitrichales Calycanthales Calycerales Campanulales Campynematales Can

ellales Cannales CapparalesCaprifoliales Cardiopteridales Carduales Caricales Carle

manniales Caryophyllales Cassiales Casuarinales CelastralesCentrolepidales Cephalot

ales Cephalotaxales Ceratophyllales Cercidiphyllales Chenopodiales Chironiales Chlor

anthalesChrysobalanales Cinchonales Circaeasterales Cistales Citrales Cocosales Col

chicales Columelliales CombretalesCommelinales Connarales Convolvulales Coriarial

es Cornales Corylales Corynocarpales Crassulales CrossosomatalesCucurbitales Cunn

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

40

inghamiales Cunoniales Cupressales Cycadales Cyclanthales Cymodoceales Cynarales

CynomorialesCyperales Cyrillales Cytinales

Daphnales Daphniphyllales Dasypogonales Datiscales Degeneriales Desfontainiales D

iapensiales Didymelales DillenialesDioales Dioncophyllales Dioscoreales Diospyrales

Dipentodontales Dipsacales Droserales

Ebenales Echiales Elaeagnales Elaeocarpales Elatinales Elodeales Emmotales Empetr

ales Ephedrales Ericales EriocaulalesErythropalaless Escalloniales Eucommiales Euph

orbiales Eupomatiales Eupteleales Euryalaless

Fabales Fagales Falcatifoliales Ficales Flacourtiales Flagellariales Fouquieriales Franc

oales Frangulales

Galiales Garryales Geissolomatales Gentianales Geraniales Gesneriales Ginkgoales G

laucidiales Globulariales GnetalesGoodeniales Greyiales Griseliniales Grossulariales

Grubbiales Gunnerales Gyrocarpales Gyrostemonales

Haemodorales Haloragales Hamamelidales Hanguanales Heisteriales Helleborales He

lwingiales Himantandrales HippuridalesHomaliales Hortensiales Huales Huerteales

Hydatellales Hydnorales Hydrangeales Hydrastidales HydrocharitalesHydropeltidales

Hydrostachyales Hypericales Hypoxidales

Icacinales Illiciales Iridales Irvingales Iteales Ixiales

Jasminales Juglandales Julianiales Juncaginales Juncales

Lacistematales Lactoridales Lamiales Lardizabalales Laurales Lecythidales Ledocarpal

es Leitneriales LentibularialesLigustrales Liliales Limnanthales Linales Liriales Loasal

es Lobeliales Loganiales Lonicerales Loranthales LowialesLythrales

Magnoliales Malpighiales Malvales Marathrales Marcgraviales Mayacales Medusagy

nales Medusandrales MelanthialesMelastomatales Meliales Melianthales Meliosmal

es Menispermales Menyanthales Metteniusales MitrastemonalesMiyoshiales Moni

miales Moringales Musales Myricales Myristicales Myrothamnales Myrsinales Myrta

les

Najadales Nandinales Narcissales Nartheciales Nelumbonales Nepenthales Neuradal

es Nitrariales Nolanales NothofagalesNyctaginales Nymphaeales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

41

Ochnales Oenotherales Olacales Oleales Onagrales Oncothecales Opuntiales Orchid

ales Oxalidales

Saxifragales Pandales Pandanales Papaverales Paracryphiales Parasitaxales Paridales

Parnassiales PassifloralesPenaeales Pennantiales Pentaphragmatales Peridiscales P

etiveriales Petrosaviales Phellinales Philydrales PhyllanthalesPhysenales Phytolaccal

es Picramniales Pinales Pinguiculales Piperales Pittosporales Plantaginales Platanale

sPlumbaginales Poales Podocarpales Podophyllales Podostemales Polemoniales Pol

ygalales Polygonales PontederialesPortulacales Posidoniales Potamogetonales Primu

lales Proteales

Quercales Quillajales Quintiniales

Rafflesiales Ranunculales Rapateales Resedales Restionales Rhabdodendrales Rhamn

ales Rhinanthales RhizophoralesRhodorales Rhoipteleales Roridulales Rosales Rouss

eales Rubiales Ruppiales Rutales

Sabiales Salicales Salvadorales Samolales Samydales Sanguisorbales Santalales Sapin

dales Sapotales SarracenialesSaxegotheales Saxifragales Scheuchzeriales Sciadopitya

les Scleranthales Scrophulariales Scyphostegiales SedalesSilenales Simmondsiales S

milacales Solanales Sphenocleales Sphenostemonales Stangeriales Stellariales Stemo

nalesStilbales Stylidiales Styracales Surianales

Taccales Tamales Tamaricales Taxales Taxodiales Tecophilaeales Terebinthales Tern

stroemiales Theales TheligonalesThymelaeales Tiliales Tofieldiales Torricelliales Tov

ariales Tribelales Trilliales Trimeniales Triuridales TrochodendralesTropaeolales Turn

erales Typhales

Ulmales Urticales

Vacciniales Vahliales Vallisneriales Velloziales Veratrales Verbenales Viburnales Vinc

ales Violales Viscales VitalesVochysiales

Welwitschiales Winterales

Xanthorrhoeales Ximeniales Xyridales

Zamiales Zingiberales Zosterales Zygophyllales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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ALPHABETICAL LISTING OF ALL FAMILY NAMES OF SEED PLANTS WITH LINKS

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In some cases clicking on a family synonym will send you directly to the subfamily in

which the genus of the basionym is to be found

Abietaceae Abolbodaceae Abrophyllaceae Acalyphaceae Acanthaceae [Acanthochla

mydaceae] Acarnaceaee AceraceaeAchariaceae Achatocarpaceae Achradaceae Achy

ranthaceae Acmopylaceae Acoraceae Acristaceae ActaeaceaeActinidiaceae Actinota

ceae Amentotaxaceae Adenogrammaceae Adoxaceae Aegialitidaceae Aegicerataceae

Aegilopaceae Aeginetiaceaee Aegiphilaceae Aesculaceae Aextoxicaceae Agapanthac

eae Agavaceae AgdestidaceaeAgialidaceae Agrimoniaceae Agrostidaceae Ailanthace

ae Aitoniaceae Aizoaceae Akaniaceae Alangiaceae AlchemillaceaeAldrovandaceae A

lismataceae Alliaceae Allioniaceae Allophylaceae Aloaceae Alopecuraceae Alpiniacea

e AlseuosmiaceaeAlsinaceae Alsodeiaceae Alstroemeriaceae Altingiaceae Alzateacea

e Amaranthaceae Amaryllidaceae Amborellaceae Ambrosiaceae Amentotaxaceae A

mmanniaceae Ammiaceae Amomaceae Ampelidopsaceae Amygdalaceae Amyridacea

eAnacardiaceae Anagallidaceae Anarthriaceae Anchusaceae Ancistrocladaceae Andr

omedaceae AndropogonaceaeAndrostachyaceae Androsynaceae Anemarrhenaceae

Anemonaceae Angelicaceae Anisophylleaceae AnnonaceaeAnomochloaceae Anopter

aceae Anrederaceae Anthemidaceae Anthericaceae Anthobolaceae Antidesmataceae

Antirrhinaceae Antoniaceae Aparinaceae Aphanopetalaceae Aphloiaceae Aphyllanth

aceae Apiaceae ApocynaceaeApodanthaceae Aponogetonaceae Aporusaceae Apose

ridaceae Apostasiaceae Aptandraceae AquifoliaceaeAquilariaceae Aquilegiaceae A

raceae Aragoaceae Araliaceae Aralidiaceaee Araucariaceae Arbutaceae Arceuthidace

aeArceuthobiaceae Arctostaphylaceae Arctotidaceae Ardisiaceae Arecaceae Argoph

yllaceae Arisaraceae AristolochiaceaeAristoteliaceae Arjonaceae Armeriaceae Artem

isiaceae Arthrotaxidaceae Artocarpaceae Arundinaceae ArundinellaceaeAsaraceae A

sclepiadaceae Ascyraceaee Asiraceae Asparagaceae Asperulaceae Asphodelaceae As

pidistraceaeeAsteliaceae Asteraceae Asteranthaceae Asteropeiaceae Astragalaceae

Astrocarpaceae AthanasiaceaeAtherospermataceae Atriplicaceae Atropaceae Aucuba

ceae Aurantiaceae Austrobaileyaceae Austrotaxaceae AvenaceaeAverrhoaceae Avet

raceae Avicenniaceae Azaleaceae Azimaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Balanitaceae Balanopaceae Balanophoraceae Balsameaceae Balsaminaceae Bambusa

ceae Barbaceniaceae BarbeuiaceaeBarbeyaceae Barclayaceae Barringtoniaceae Base

llaceae Bataceae Baueraceae Bauhiniaceae Baxteriaceae BegoniaceaeBehniaceae B

elangeraceae Belloniaceae Bembiciaceae Berberidaceae Berberidopsidaceae Berryac

eaee BersamaceaeBertyaceae Berzeliaceae Besleriaceae Betaceae Betulaceae Biebe

rsteiniaceae Bifariaceae Bignoniaceae BischofiaceaeBixaceae Blakeaceae Blandfordia

ceae Blattiaceae Blepharocaryaceae Blitaceae Blyxaceae Boerlagellaceae Bolivaracea

eBombacaceae Bonnetiaceae Bontiaceae Boopidaceae Boraginaceae Borassaceae B

oroniaceae BoryaceaeBotryodendraceae Bougainvilleaceae Boweniaceae Brassicacea

e Brachycaulaceae Bretschneideraceae BrexiaceaeBromeliaceae Brownlowiaceae Br

unelliaceae Bruniaceae Brunoniaceae Brunsvigiaceae Bryoniaceae BuchneraceaeBuci

daceae Buddlejaceae Buglossaceae Bulbocodiaceae Bumeliaceae Bupleuraceae Burc

hardiaceae BurmanniaceaeBurseraceae Butneriaceae Butomaceae Buxaceae Byblida

ceae Byttneriaceae

Cabombaceae Cacaoaceae Cactaceae Caesalpiniaceae Caladiaceae Calamaceae Calce

olariaceae CalectasiaceaeCalendulaceae Callaceae Callicomaceae Calligonaceae Calli

traceae Callitrichaceae Calochortaceae CalophyllaceaeCalthaceae Calycanthaceae Ca

lyceraceae Cambogiaceae Camelliaceae Campanulaceae CampynemataceaeCanacom

yricaceae Canellaceae Cannabaceae Cannaceae Canopodaceae Canotiaceae Cansjera

ceae CapparaceaeCaprariaceae Caprifoliaceae Cardiopteridaceae Carduaceae Carica

ceae Carissaceae Carlemanniaceae CarpinaceaeCarpodetaceae Cartonemataceae Ca

ryocaraceae Caryophyllaceae Casearieaceae Cassiaceae CassipoureaceaeCassythacea

e Castaneaceae Castelaceae Casuarinaceae Catesbaeaceae Cathedraceae Cecropiace

ae CedraceaeCedrelaceae Celastraceae Celosiaceae Celtidaceae Centaureaceae Cen

trolepidaceae Centroplacaceae CepaceaeCephalanthaceae Cephalotaceae Cephalota

xaceae Cerastiaceae Ceratoniaceae Ceratophyllaceae CerberaceaeCercidiphyllaceae

Cercocarpaceae Cercodiaceae Cereaceae Cerinthaceae Ceroxylaceae Cestraceae Cev

alliaceaeChailletiaceae Chamaedoraceae Chamaemoraceae Chamelauciaceae Chaun

ochitonaceae Chelidoniaceae ChelonaceaeChenopodiaceae Chimonanthaceae Chingi

thamnaceae Chionographidaceae Chiranthodendraceae ChironiaceaeChlaenaceae Ch

loanthaceae Chloranthaceae Chloridaceae Chlorogalaceae Chrysobalanaceae Cicerac

eae CichoriaceaeCimicifugaceae Cinchoniaceae Circaeaceae Circaeasteraceae Cissac

eae Cistaceae Citraceae Clematidaceae CleomaceaeClethraceae Clusiaceae Cneorac

eae Cnestidaceae Cnicaceae Cobaeaceae Cochlospermaceae Cocosaceae Coffeaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Colchicaceae Coleogynaceae Columelliaceae Combretaceae Commelinaceae Comocl

adiaceae Compositae CompsoaceaeConantheraceae Connaraceae Conostylidaceae C

onvallariaceae Convolvulaceae Cordiaceae CoreopsidaceaeCoriandraceae Coriariacea

e Coridaceae Corispermaceae Cornaceae Corokiaceae Coronillaceae Correaceae Cor

rigiolaceaeCorsiaceae Corydalaceae Corylaceae Corynocarpaceae Coryphaceae Cost

aceae Cotyledonaceae CoulaceaeCoutariaceae Coutoubiaceae Crassulaceae Crescen

tiaceae Cressaceae Crinaceae Crocaceae CroomiaceaeCrossosomataceae Crotonacea

e Cruciferae Cryptaceae Crypteroniaceae Cryptocorynaceae CryptomeriaceaeCtenol

ophonaceae Cucurbitaceae Cunninghamiaceae Cunoniaceae Cupressaceae Curcumac

eae Curtisiaceae CuscutaceaeCyananthaceae Cyanastraceae Cynanchaceae Cyanella

ceae Cycadaceae Cyclanthaceae CyclantheraceaeCyclocheilaceae Cydoniaceae Cymo

doceaceae Cynaraceae Cynocrambaceae Cynomoriaceae Cyperaceae CyphiaceaeCyp

hocarpaceae Cypripediaceae Cyrillaceae Cyrtandraceae Cyrtanthaceae Cytinaceae

Dacrycarpaceae Dactylanthaceae Damasoniaceae Daphnaceae Daphniphyllaceae Das

ypogonaceae DatiscaceaeDaturaceae Daucaceae Davidiaceae Davidsoniaceae Decais

neaceae Deeringiaceae Degeneriaceae DelphiniaceaeDendrophthoaceae Columelliac

eae Detariaceae Dialypetalanthaceae Dianellaceae Dianthaceae DiapensiaceaeDicha

petalaceae Dichondraceae Diclidantheraceae Dicrastylidiaceae Dictamnaceae Didiere

aceae DidymelaceaeDidymocarpaceae Diegodendraceae Diervillaceae Digitalidaceae

e Dilatridaceae Dilleniaceae Dioaceae DionaeaceaeDioncophyllaceae Dioscoreaceae

Diosmaceae Diospyraceae Dipentodontaceae Diphylleiaceae Diplolaenaceae Dipsaca

ceaeDipterocarpaceae Dirachmaceae Disanthaceae Diselmaceae Dodonaeaceae Do

mbeyaceae Donatiaceae DorsteniaceaeDortmannaceae Doryanthaceae Drabaceae D

racaenaceae Dracontiaceae Drimyidaceae Droseraceae DrosophyllaceaeDryadaceae

Duabangaceae Duckeodendraceae Dulongiaceae Durantaceae Durionaceae Dysphani

aceae

Ebenaceae Eccremidaceae Ecdeiocoleaceae Echiaceae Echinopaceae Ehretiaceae Ela

eagnaceae ElaeocarpaceaeElegiaceae Elatinaceae Ellisiaceae Ellisiophyllaceae Elodea

ceae Elytranthaceae Embeliaceae Emblingiaceae EmmotaceaeEmpetraceae Encepha

lartaceae Engelhardtiaceae Enhalaceae Epacridaceae Ephedraceae Ephemeranthacea

eEpilobiaceae Epimediaceae Eragrostidaceae Eremolepidaceae Eremosynaceae Erica

ceae Erinaceae EriocaulaceaeEriogonaceae Eriospermaceae Erodiaceae Erycibaceae

Eryngiaceae Erysimaceae Erythroniaceae ErythropalaceaeErythrospermaceae Erythro

xylaceae Escalloniaceae Eschscholziaceae Eucomidaceae Eucommiaceae Eucryphiace

aeEuonymaceae Eupatoriaceae Euphorbiaceae Euphrasiaceae Euphroniaceae Eupo

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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matiaceae Eupteleaceae EuryalaceaeEustrephaceae Euthemidaceae Eutocaceae Exb

ucklandiaceae Exocarpaceae

Fabaceae Fagaceae Falcatifoliaceae Ferulaceae Festucaceae Ficaceae Ficoidaceae Fit

zroyaceae FlacourtiaceaeFlagellariaceae Flindersiaceae Foetidiaceae Forestieraceae

Fothergillaceae Fouquieriaceae Fragariaceae FrancoaceaeFrangulaceae Frankeniacea

e Fraxinaceae Fraxinellaceae Freycinetiaceae Fritillariaceae Fuchsiaceae Fumariacea

eFunkiaceae

Gaiadendraceae Galacaceae Galanthaceae Galaxiaceae Galedupaceae Galeniaceae G

aliaceae Garciniaceae GardeniaceaeGardneriaceae Garryaceae Geissolomataceae Ge

itonoplesiaceae Gelsemiaceae Geniostomaceae GentianaceaeGeonomataceae Geosir

idaceae Geraniaceae Gerrardinaceae Gesneriaceae Gethyllidaceae Gilliesiaceae Gina

lloaceaeGinkgoaceae Gisekiaceae Gladiolaceae Glaucidiaceae Glechomaceae Glinace

ae Globulariaceae Gnaphaliaceae GnetaceaeGoetzeaceae Gomortegaceae Gomphiac

eae Gomphrenaceae Gonystylaceae Goodeniaceae Gordoniaceae GouaniaceaeGoup

iaceae Gramineae Gratiolaceae Grewiaceae Greyiaceae Grielaceae Griseliniaceae Gr

onoviaceae GrossulariaceaeGrubbiaceae Guaiacanaceae Guamatelaceae Guettardac

eae Gunneraceae Gustaviaceae Guttiferae GyrocarpaceaeGyrostemonaceae

Hachetteaceae Haemanthaceae Haemodoraceae Halesiaceae Hallieraceae Halocarpa

ceae Halophilaceae HalophytaceaeHaloragaceae Hamamelidaceae Hanguanaceae H

aptanthaceae Harmandaceae Hebenstretiaceae HectorellaceaeHederaceae Hedyosm

aceae Hedyotidaceae Hedysaraceae Heisteriaceae Heleniaceae Heliamphoraceae H

elianthaceaeHelianthemaceae Helichrysaceae Heliconiaceae Helicteraceae Heliotropi

aceae Helleboraceae Heloniadaceae HelosidaceaeHelwingiaceae Hemerocallidaceae

Hemimeridaceae Hemiphylacaceae Henriqueziaceae Henslowiaceae Hermanniaceae

Hernandiaceae Herniariaceae Herreriaceae Hesperocallidaceae Heterantheraceae He

teropyxidaceae HeterostylaceaeHibbertiaceae Hibiscaceae Hilleriaceae Himantandra

ceae Hippocastanaceae Hippocrateaceae HippomanaceaeHippophaeaceae Hippurida

ceae Hirtellaceae Holacanthaceae Homaliaceae Hopkinsiaceae Hoplestigmataceae H

ordeaceaeHornschuchiaceae Hortensiaceae Hortoniaceae Hostaceae Houstoniaceae

Huaceae Huerteaceae HugoniaceaeHumbertiaceae Humiriaceae Hyacinthaceae Hyd

atellaceae Hydnoraceae Hydrangeaceae Hydrastidaceae HydrillaceaeHydrocharitacea

e Hydrocotylaceae Hydrogetonaceae Hydroleaceae Hydropeltidaceae Hydrophylacac

eae HydrophyllaceaeHydrostachyaceae Hymenocardiaceae Hypecoaceae Hyperanth

eraceae Hypericaceae Hypopityaceae HypoxidaceaeHypseocharitaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta

ceae ImperatoriaceaeInocarpaceae Inulaceae Ionidiaceae Iriarteaceae Iridaceae Irvi

ngiaceae Isnardiaceae Isophysidaceae IteaceaeStrasburgeriaceae Ixiaceae Ixioliriace

ae Ixonanthaceae

Jalapaceae Jamboliferaceae Japonoliriaceae Jasionaceae Jasminiaceae Johnsoniaceae

Joinvilleaceae JuglandaceaeJulianiaceae Juncaceae Juncaginaceae Juniperaceae Jus

siaeaceae Justiciaceae

Kadsuraceae Kaliphoraceae Kalmiaceae Kaniaceae Kiggelariaceae Kingdoniaceae Kin

giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae

Krameriaceae

Labiatae Lacandoniaceae Lachenaliaceae Lacistemataceae Lactoridaceae Lactucaceae

Lagerstroemiaceaee LamiaceaeLampsanaceae Lanariaceae Langsdorffiaceae Lantan

aceae Lapageriaceae Lardizabalaceae Lasiopetalaceae LauraceaeLawsoniaceae Laxm

anniaceae Lecythidaceae Ledaceae Ledocarpaceae Leeaceae Leguminosae Leitneriac

eae LemnaceaeLennoaceae Lentibulariaceae Lentiscaceae Leoniaceae Leonticaceae

Lepidobotryaceae LepidocarpaceaeLepidocaryaceae Lepidocerataceae Leptaulaceae

Leptospermaceae Lepturaceae Lepuropetalaceae LeuctenbergiaceaeLibocedraceae

Licaniaceae Ligustraceae Lilaeaceae Liliaceae Limeaceae Limnanthaceae Limnocharit

aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace

ae Linderniaceae Lindleyaceae Linnaeaceae LippayaceaeLiriaceae Liriodendraceae Li

ssocarpaceae Littorellaceae Loasaceae Lobeliaceaee Loganiaceae LomandraceaeLoni

ceraceae Lopeziaceae Lophiocarpaceae Lophiolaceae Lophiraceae Lophophytaceae L

ophopyxidaceae LoranthaceaeLotaceae Lowiaceae Lupulaceaee Luxemburgiaceae Lu

zuriagaceae Lychnidaceae Lyciaceae LyginiaceaeLygodisodeaceae Lysimachiaceae Ly

thraceae

Macarisiaceae Mackinlayaceae Madiaceae Maesaceae Magnoliaceae Malaceae Male

sherbiaceae MalortieaceaeMalpighiaceae Malvaceae Mangiaceae Manicariaceae Ma

paniaceae Marantaceae Marathraceae MarcgraviaceaeMartyniaceae Mastixiaceae

Matricariaceae Maundiaceae Mayacaceae Medeolaceae Medusagynaceae Peridiscac

eaeMelaleucaceae Melampyraceae Melanophyllaceae Melanthiaceae Melastomatac

eae Meliaceae Melianthaceae MelicaceaeMeliosmaceae Melittidaceae Melochiacea

e Memecylaceae Mendonciaceae Menispermaceae Menthaceae MenyanthaceaeMe

nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

Nypaceae Nyssaceae

Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

ceae Papilionaceae Pappophoraceae PapyraceaeParacryphiaceae Parasitaxaceae Par

ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

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cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

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ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 19: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

19

problem is the use of one or a very few characters that have been weighted a priori to

structure classifications (see also Garciacutea et al 2009) although it is perhaps not

surprising that such an approach sometimes results in the delineation of taxa that have

indeed turned out to be quite distinct (cf eg Burtt Astragalus versus Oxytropis)

Taxa recognized here are monophyletic yet many are polythetic at the

morphological level that is they lack unique features characterizing (defining) all and

only members of that family they can be recognised phenetically only by the unique

combinations of characters that they posess This is the result of evolution any

synapomorphy characterizing a taxon may be lost or modified beyond easy recognition

in some of its members or the synapomorphy may appear to be identical to a feature

that has evolved in parallel in a quite unrelated plant That plant groups are polythetic is

almost as much a feature of monophyletic taxa as the paraphyletic taxa common in

evolutionary classifications Some families now include substantial variation as

phenetically distinct derived groups are placed in their proper phylogenetic position -

examples are Ericaceae which include the erstwhile Empetraceae a wind-pollinated

group that is florally very different and the overwhelmingly large-flowered and animal-

pollinated Plantaginaceae which now include several derived small-flowered aquatic

and wind-pollinated groups that were previously placed in separate families

Thus some families as delimited here may not be easy to recognize However

remember that detecting relationships - use whatever characters you can even if they

are not obvious - and naming a plant - focus on easy-to-see characters that may not

reflect relationships - are quite different problems Taxa although natural may not be

readily recognizable indeed it was in exactly this context that Lamarck worked out the

basic principles of writing dichotomous keys in 1778 (see Scharf 2007 for keys and the

like) Of course Lamarcks idea of nature was very different from ours - he thought that

there was some kind of continuum of form on which living organisms were to be

situated with no real gaps anywhere - but this meant that his genera (for example)

might well not be sharply distinct from each other even if each were part of the real

continuum that was life Lamarcks separation of the establishment of relationships and

groupings in a system and identification of the plants in that system have been widely

accepted (but cf eg Godfray amp Knapp 2004 many of the contributors to Heywood et

al 2007) Perhaps the best way of identifying plants at the family level is by well-made

multiple access keys as in Watson and Dallwitz (1992a onwards family limits there may

differ substantially from those adopted here) Multiple access keys free users from the

constraints of dichotomous keys in which particular characters are needed at each step

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

20

of the identification process before there can be further progress Instead those

characters that happen to be evident on a specimen can be used in whatever order is

convenient when linked to illustrations glossaries etc their power is enormous (see

Dallwitz et al 2000 [2006] for the principles underlying their construction and use)

Nevertheless dichotomous keys such as those of Hutchinson (1973) and Franz Thonner

(Geesink et al 1981) have their uses Of course taxa coming out adjacent in keys may

well not be at all related

(When identifying large numbers of plants even more efficient than either style of

identification and certainly lots more fun is sight identification Unless you have a

photographic memory you have to build up your knowledge of comparative plant

morphology - on which the ability to make accurate identifications depends - by

repeated observation When faced with an unknown plant I always look for leaf teeth

and stipules distinctive hair types especially stellate and T-shaped hairs glands and

punctations of any sort the presence of latex or other exudate and check leaf insertion

smelling crushed leaves can also be helpful In this context nodal anatomy can usually

be checked using a razor and a hand lens or even simply carefully examining leaf scars

The short paragraphs added after most families may help in confirming familial

identifications)

For the record and for the little that it is worth there are 4 orders and 13 families

of gymnosperms characterised on these pages and together they include some 82

genera and 947 species For angiosperms comparable figures are 56 orders 445

families 13208 genera and 261750 species (of which monocots include 11 orders 89

families 2759 genera and 52760 species) Note however that higher mathematics

was never my strong point and anyway these are pretty meaningless figures even for

species which many (but not all) might concede smacked slightly more of reality than

other taxa estimates range as high as 422000 (Govaerts 2001) Furthermore numbers

of genera and species change daily Nevertheless as emphasized here families are

useful in teaching we as a community can ensure that their limits remain largely stable

and by concentrating on relatively few of them one can gain some familiarity with much

of the worlds flora For summaries of the sizes of orders and families and of the general

arrangement followed in these pages see also the Statistics page itself

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

21

A PHYLOGENETIC TREE OF SEED PLANTS (THE MAIN TREE)

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

22

The Main Tree above is a conservative summary of well supported relationships

between the major clades of extant seed plants Reading - interpreting and

understanding - such a phylogenetic tree is not necessarily easy so reading a tree may

require a bit of practice OHara (eg 1988 1992) provides an accessible introduction to

thinking and talking about history in the context of looking at phylogenetic trees More

recently Krell and Cranston (2004) Crisp and Cook (2005) and others have emphasized

how careful one must be when interpreting and talking about evolution in ladderized

trees in particular and phylogenetic trees in general Gregory (2008) provides a

particularly detailed discussion of this problem In ladderized trees the smaller (in terms

of numbers of terminals) sister taxon is consistently shown on the same side at every

node thus the trees tend to be pectinate like the teeth of a comb however the

horizontal axis that is as a result so evident has no polarity or particular direction The

use of the adjective basal when discussing phylogenetic trees is especially dangerous

(see also Wojciechowski et al 2004 D Soltis et al 2005b) When I use the term and the

context is not otherwise clear I am referring to the pectinations at the base of a

ladderised tree Remember that when talking about sister taxa one can never be basal

to the other or older or younger although members of one can be more derived - in the

sense of having more apomorphies - than the other (but even then one has to be

careful) The words primitive and advanced and lower and higher should

practically never be used especially when talking about taxa Amborellaceae and

Pinaceae are sister to all other flowering plants and Pinales and are basal to the crown

groups of all other flowering plantsPinales respectively but that does not mean they

are thereby primitive Pinaceae in particular have numerous apomorphies The word

plesiomorphic is far less loaded than primitive and can be used to talk about

individual characters

Most trees here have been more or less ladderized as is the one above The

ladderization may be imperfect for example you can see that on the Main Tree

asterids with ten orders follow rosids and relatives which have seventeen although

this is in part because of the currently unresolved nature of relationships between the

main clades of core eudicots (this is likely to change soon) The Main Tree could have

been drawn with Amborella Acorus or a host of other taxa at the far right without

offending any relationships phylogenetic trees are like mobiles the only fixed points

being the nodes Despite the problems just mentioned pectination interpreted

carefully has its value and it is not an arbitrary process As one reads the terminals of a

pectinate tree from left to right adjacent terminals will be separated by apomorphies

that will cumulate in an appropriate fashion When thinking of a book or a herbarium

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

23

sequence (see Haston et al 2007) this is of some value Nymphaeales and

Austrobaileyales are here adjacent on the tree but they could be separated by

hundreds of families in the sequence merely by rotating only the node from which

Nymphaeales and their sister group (all other angiosperms minus about 75 species)

arise If adjacent in a book or herbarium or as in these summary pages then it is

relatively easy to relate their characters both apomorphic and plesiomorphic but if

separated by hundreds of pages or two floors in a large building then it is less easy to

get anything from the sequence (Since all orders in the pages of the main part of this

site are preceded by the apomorphies of all nodes immediately below them in the seed

plant phylogeny and because of the linkages that have been built in to the site this

problem is much less serious there there is no sequence) So the particular sequence of

all seed plants that is based on a phylogenetic classification but is to be used in herbaria

or any other place where taxa have to be arranged linearly as in this part of APweb

will be one that maximizes the number of taxa that are both successive branches of the

tree and placed successively in the sequence Since specimens are generally filed under

families the outline of a new family sequence for arranging herbaria and books can now

be suggested (Haston et al 2007) and such a standardized sequence will further help

teaching and learning about plants

ON THINKING ABOUT APOMORPHIES

Identifying apomorphies is important because understanding the evolution of

morphology in the broad sense - ie including anatomy chemistry etc - in the context

of diversification is one of our major goals For identifying apomorphies several

preconditions must be met One needs to have an accurate well supported phylogeny

one has to have examined the right taxa both from the point of view of morphology and

molecules one has to have coded the characters right (ie delimited states

appropriately) and one has to use an appropriate model of evolution when fitting the

variation to the tree (see eg Omland 1999 Stevens 2006b) If the distributions of

apomorphies below are compared with those in other studies (eg Turgeon et al 2001

Bremer et al 2001 Endress 2001 Albach et al 2001a Judd amp Olmstead 2004 D Soltis

et al 2005b Zhang et al 2006 Judd et al 2007) the reader may find a number of

differences Although I have integrated such studies as far as is possible there are five

reasons that link to the preconditions just mentioned why there may be differences

where features are placed on trees

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

24

1 I may not have found all the information about a particular character there may be

disagreement over its interpretation or I have added information

2 The sampling of nearly all molecular studies is very incomplete (see Salisbury amp Kim

2001 for problems caused by sampling) indeed it is commonplace to decry the

incompleteness of molecular sampling But not only is the sampling in molecular studies

often less than we might wish that of the morphological and chemical characters whose

evolution we are interested in understanding is also often very poor So for many

anatomical chemical and embryological characters that are confidently said to

characterise families and other groups we all too often have no idea if those characters

are applicable to the whole clade or just to a subgroup within that clade However as

our sampling improves we can locate changes on the tree more precisely if in

unexpected positions Thus is is sometimes suggested that Ericaceae have ellagic acid

(Soltis et al 2005b) in fact the little evidence we have indicates that only a small clade

of some 80 species has ellagic acid and the rest of the family - itself alone about one

third of all Ericales - does not Similarly Ericaceae are noted for their

ectendomycorrhizal associations but Enkianthus sister to all other Ericaceae

apparently lacks these (Abe 2005) Thus these mycorrhizae and their associated

distinctive hair roots are unlikely to be apomorphies of Ericaceae - however it appears

that other families near Ericaceae may also have ectendomycorrhizae (see Asai 1934)

3 I am fitting characters to a very conservative tree with many polytomies although the

nodes that are utilised are for the most part strongly supported polytomies make the

optimisation of characters that is the assigment of character state change to a

particular node on the tree notably difficult (eg Madison amp Madison 2002) In nearly

all studies of the evolution of characters distributions of characters are optimised on a

more or less fully resolved tree and the construction of supertrees may yield yet more

detailed hypotheses of relationships (for literature on supertrees see Cotton amp

Wilkinson 2007 2008) Of course some nodes on such fully resolved trees andor

supertrees may have little support and optimisations of characters on such trees may

carry correspondingly little conviction Even parts of some of the trees used here have

poor support eg relationships within aquatic Alismatales etc although I indicate

when this is the case

4 Exactly how one goes about optimising a character on a tree is critically important

Even using simple parsimony optimisations (ACCTRAN or DELTRAN ACCelerated

TRANsitions or DELayed TRANsitions) the position of synapomorphies on trees - and

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

25

hence our ideas of evolution - may differ and this is still more true if one uses maximum

likelihood or Bayesian methods and makes apparently reasonable suggestions about

weighting gains over losses (eg Donoghue amp Ackerley 1996 Cunningham et al 1998

Omland 1997 1999 Ree amp Donoghue 1999 Polly 2001 Webster amp Purvis 2001

Ronquist 2004 Crisp amp Cook 2005) Sannier et al (2007) show how in in Arecaceae that

where on a phylogenetic tree one might peg changes in microsporogenesis will depend

on the methods one uses to do this and Pedersen et al (2007) discuss the sometimes

very substantial effect of node support on the posterior probabilities of ancestral

character states Here I use parsimony optimization not always as explicit as it might be

but I have often indicated where there are particularly important uncertainties as to the

positions of particular character changes on the tree

5 Finally although I have paid quite a lot of attention to the delimitation of the

character states that make up all the characterizations I have not spent enough time on

this critical operation If we are interested in understanding evolution then fitting the

basic variation - not character states - to a tree in principle allows greater flexibility in

understanding morphology in the context of local phylogenies (see also Stevens 2000

Endress 2005c) However many character states used here are delimited globally that

is they are circumscribed in the context of the variation shown by individual characters

across all angiosperms andor in the context of classic ideas of character evolution

Character states often have arbitrary limits and serve best to communicate

information whether they are in fact suitable for either phylogenetic analysis or

understanding evolution are separate issues Studies have rather unsurprisingly

perhaps but importantly shown that dividing the one character into different sets of

states may yield differing ideas of evolution of that character (eg Lamb Frye amp Kron

2003 Hibbett 2004) When looking at trees on which character states are optimised

one should bear in mind the problems surrounding the delimitation of states (eg

Stevens 2000 2006b) and the danger of using pollen or other types - constructs

based on many characters that vary independently but which effectively get lost in

these types thus Blackmore et al (2009) decomposed the pollen types in Asteraceae

into 52 characters

I use Remanes three main criteria of homology or better similarity when

determining the basic similarity of structures on different organisms (see Remane 1952)

These criteria are special properties position and intermediates Special properties

include anatomical or chemical characters gene expression data etc Position refers

to the position of an organ with respect to landmarks on the plant Although plants are

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

26

plants and landmarks are not as fixed as one might like trying to work out the relative

positions of parts is a good way of understanding morphology so long as one does not

try to out-do Procrustes Intermediates include intermediates found on the same

plant or on different plants Intermediates may observed during development or be

adult structures Thus one may find changes during development which suggest what

the nature of a particular structure is and very different stuctures on different plants

may turn out to be similar early in development Or one can simply compare different

taxa and see that structures that appear to be very different when comparing two

different taxa can be linked morphologically by looking at other taxa

Nevertheless the use of these criteria may not yield an unambiguous answer as to

what a structure is even given a solid phylogeny and an improved understanding of

development (see Jaramillo amp Kramer 2007 for a useful discussion) As Endress (2005c)

observed a number of features - position function development shape anatomy

histology gene activity and relationships to other taxa that clearly have petals - can be

used to distinguish a petal (for example) from other floral structures if a petal does not

have one of these features is it thereby not a petal Thus Maturen et al (2005) recently

found that floral organ diversity genes (B and C) were expressed in the large white

inflorescence bracts of Cornus (see also Costa et al 2005) Peney et al (2005) noted that

not all monosulcate pollen grains in monocots have the same developmental pathway

and that as a result such pollen might not have the same ancestral state Reeves and

Olmstead (2003) suggested that the genetic mechanisms causing monosymmetry in

Lamiales and Solanales were different and Serna and Martin (2006) described similar

problems with the development of hairs in Arabidopsis when compared with that of

hairs in Antirrhinum and Solanaceae Indeed as one perhaps might expect delimitation

of states and characters does not necessarily become easier with increasing knowledge

of development etc Thus Buzgo et al (2004) Matthews and Endress (2005) and others

have shown how hard it can be to distinguish between eg prophylls and other floral

structures as their behaviour is studied during the course of the development of the

flower Similarly at what concentration is a particular secondary metabolite deemed to

be present (Waterman (2007)

The validity of the approach used here that of fitting morphological variation to a

largely molecular-based tree may be questioned However I think it rather unlikely that

well-supported molecular branches will be overturned by morphological data Indeed

analyses of morphological data alone do provide support for many of the clades evident

in molecule-only analyses and analyses with morphological and molecular data

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

27

together may lead to increased support for clades (eg Hufford 1992 Nandi et al 1998

[but adding morphological data reduces support for a number of critical clades too]

Doyle amp Endress 2000 however in none of these papers is the use of morphology

without ambiguity) It is unfortunately clear that the use of morphology alone may not

suggest problems in the phylogenetic placement of taxa that later turn out to have been

wrongly included (eg Zhang et al 1992) Thus I have been wary of putting much weight

on clades that have only morphological support but note that I have treated molecule-

based clades with low bootstrap or jacknife support values (esp below 70) or low

posterior probabilies (below 095) likewise Although I may have been mistaken in

placing so much emphasis on molecular data in terms of providing the basic

phylogenetic framework for angiosperms morphological and molecular data are only

very rarely in irreconcilably strong conflict There are indeed a few places where the

conflict seems extreme These include the relative positions of the Monimiaceae and

Hernandiaceae (Laurales) the position of Hanguanaceae (Commelinales [as here] or

Zingiberales) and of Triplostegia (is it in Dipsacaceae or Valerianaceae - see

Dipsacales) Fossils are unlikely to affect the topologies of the trees presented here but

see below for their importance in understanding morphological evolution in general and

the evolution of angiosperms in particular Nevertheless some largely reject the idea

that trees based on molecular data alone can recover phylogenetic relationships

especially when branching points are old and prefer to used trees based on analysis of

morphological data including those taken from fossils (Hilton amp Bateman 2006 Farjon

2007)

All in all however the extent of the congruence between morphological and

molecular data is impressive and heartening and many clades can be characterised

morphologically It seemed in 1998 that there were no unambiguous morphological

synapomorphies for angiosperm orders (K Bremer 2000) and this is still true if by

unambiguous is meant non-homoplasious However many orders and other clades

have synapomorphies even if these may be indistinguishable at least at the current

level of morphological and developmental knowledge from parallel occurences

elsewhere As our morphological knowledge increases so too does the number of

apomorphies

Endress and Matthews (2006a) emphasize the importance of tendencies and

developmental constraints when thinking about characters of clades within the rosids -

many characters or character combinations occur in a rather sporadic fashion within

the clade and apparently notably less frequently outside it There are a number of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

28

examples of tendencies here and in the individual order pages such as the features

enclosed by parentheses in groups above the level of family eg see (cuticular waxes

as aggregated rodlets) for the commelinids and the discussion of the distribution of

polyandry that is flowers with numerous stamens in the asterid I + II groups etc

Tendencies may involve more than single characters When thinking of the

associations of features like integument thickness and vascularization of the integument

(Endress amp Matthews 2006a) one can imagine fairly simple developmental

preconditions being operative As Endress and Matthews (2006a) note it is difficult to

imagine vascular tissue developing in an integument only two cells thick Changes here

would be loosely correlated if morphologically linked However the strongly correlated

changes noted by Givnish et al (2005) are ecologically linked but are presumably

morphologicallydevelopmentally independent When there seem to be characters

evolving more or less together tests can be carried out to see if the changes are

concentrated on certain branches of the tree (eg Maddison 1990 Sanderson 1991

Maddison amp Maddison 2000) Such tests have rarely been carried out at higher levels in

angiosperms

One of the most striking examples of a tendency is the distribution of N-fixation

restricted as it is to a monophyletic group of four clades although it has arisen seven (or

perhaps several more) times independently within the N-fixing clade and several

members of two quite different kinds of bacteria are involved (eg Clawson et al 2004

Elliott et al 2007 Sprent amp James 2007) Other examples of work that bears on the issue

of tendencies include the findings that flowers of polysymmetrical Arabidopsis have

genes like TCP1 that are expressed asymmetrically during early development and TCP1

is a probable orthologue of the well-known CYC gene of Antirrhinum that is involved in

the development of monosymmetric slowers there (Cubas et al 2001 Costa et al 2005

etc) Parallelism might build on this underlying morphologically cryptic monosymmetry

even if details of the genetic mechanisms causing the monosymmetry evident in

particular groups may be different (Reeves amp Olmstead 2003 Cubas 2004 see above)

Similarly there have been several recent suggestions that the capability to synthesise a

particular metabolite may be switched off but not lost and so can sometimes be

reacquired (eg Wink amp Witte 1983 Wink 2003 Liscombe et al 2005 Larsson 2007

Waterman 2007) Hence perhaps the rather spotty distribution of many secondary

metabolites like ellagic acid the indole alkaloid camptothecin iridoids etc when

considered in the context of phylogenies As a non-botanical example - but a rather nice

one - Salwini-Plawen and Mayr (1961) suggested some time ago that there has been

considerable parallelism (40-65 or more independent origins) in the evolution of eyes in

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

29

metazoans However Pax 6 seems to be a master control gene that is involved in eye

formation perhaps throughout all bilateralians (eg Gehring amp Ikeo 1999 Erwin amp

Davidson 2002) which changes the nature of the problem Note however that in

general our invocation of evolutionary tendencies to explain the patterns of

distributions of characters is really an admission that we do not understand those

patterns

As we find out more about variation we will probably find fewer and fewer features

constant throughout a group Most unqualified statements of presence and absence

should properly be qualified as usually present or usually absent if one is thinking of

the characterisations on the order pages as encompassing the total variation within a

clade Thus Pistia alone among monocots as so far known has sieve tube plastids with

starch grains not protein crystals However this does not affect the fact that sieve tube

plastids with cuneate protein crystals and lacking starch grains are an apomorphy for

monocots Furthermore fossils in a number of cases suggest character combinations

unknown in extant taxa as may be seen in the discussions of Fagaceae Platanaceae

Iteaceae Calycanthaceae etc Confusing the issue there may be questions as to where

exactly on the tree a particular fossil is to be placed (eg see Nymphaeaceae

Calycanthaceae Archaefructus etc)

To summarize given our current understandings of both phylogenies and

characters evolution of some characters in which we are interested seems very labile

(see eg D Soltis et al 2005b Endress and Matthews 2006a Stevens 2006b Ekman et

al 2008 for a good recent study) and I have been cautious when talking about character

evolution Much effort must continue to be spent in summarizing characters of well-

established clades at all levels providing features by which they may be recognized and

signaling synapomorphies Remember that (1) the basic morphological anatomical and

chemical knowledge of many critical taxa is woefully incomplete (2) different

assumptions about character evolution may greatly affect the position of

synapomorphies on trees (3) in many cases relationships within and between many

groups are too uncertain at present to worry very much about synapomorphies and

(4) we must be clear about what we do and do not not know As mentioned above I

have indicated in a number of places where there are particularly important

uncertainties as to where characters should be placed on the tree Nevertheless it is a

relatively easy matter to update notes such as these and it can be a simple matter to

incorporate new data on characters that have never before been considered in the

context of a tree Much basic - and unfortunately perhaps unfashionable - work must be

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

30

carried to clarify the distribution of morphological anatomical and chemical characters

for examples of what can to be done see the work of P K Endress and collaborators

(floral morphology and development) and S R Jensen and collaborators (iridoids) But

acquiring information about nodal anatomy cell and tissue distribution nodal anatomy

and the like is not difficult at all and should be generally encouraged

But all this is merely a necesary prelude to the understanding of evolution For this

one needs to know a lot more including dating the phylogeny understanding the

function(s) of characters factoring in relevant aspects of the palaeoenvironment in

which an apomorphy first appeared etc

SUMMARY OF THE SYSTEM

Below is a formalised summary of the relationships within orders of the families of seed

plants There are a few families that are not recognised even as options in APG II and

vice versa as well as a few extra orders The families to be recognised in the most

recent edition of Mabberleys The Plant Book (Mabberley 2008) are also largely

consistent with those below But all differences are trivial and will - I hope - eventually

disappear

Square brackets - [] - enclose clades the plus sign - + - designates sister taxa a

comma - - denotes part of a polytomy and quotation marks - - denotes a

paraphyletic group

SEED PLANTS

GYMNOSPERMS

Cycadales

Cycadaceae + Zamiaceae

Ginkgoales

Ginkgoaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

31

Pinales

Pinaceae [[Araucariaceae [Podocarpaceae + Phyllocladaceae]] [Sciadopityaceae

[Taxaceae + Cupressaceae]]]

Gnetales

Ephedraceae [Gnetaceae + Welwitschiaceae]

ANGIOSPERMSFLOWERING PLANTS

Amborellales

Amborellaceae

Nymphaeales

Hydatellaceae [Cabombaceae + Nymphaeaceae]

Austrobaileyales

Austrobaileyaceae [Schisandraceae + Trimeniaceae]

Chloranthales

Chloranthaceae

MAGNOLIIDS

Magnoliales

Myristicaceae [Magnoliaceae [[Himantandraceae + Degeneriaceae] [Eupomatiaceae +

Annonaceae]]]

Laurales

Calycanthaceae [[Siparunaceae [Gomortegaceae + Atherospermataceae]] [Monimiaceae

[Hernandiaceae + Lauraceae]]]

Canellales

Canellaceae + Winteraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

32

Piperales

[Hydnoraceae + Aristolochiaceae] [Piperaceae + Saururaceae]

MONOCOTS

Acorales

Acoraceae

Alismatales

Araceae [Tofieldiaceae [[Alismataceae [Hydrocharitaceae + Butomaceae]]

[Scheuchzeriaceae [Aponogetonaceae [Juncaginaceae [Maundiaceae [[Posidoniaceae

[Ruppiaceae + Cymodoceaceae]] [Zosteraceae + Potamogetonaceae]]]]]]]

Petrosaviales

Petrosaviaceae

Dioscoreales

Nartheciaceae [[Taccaceae + Thismiaceae] [Burmanniaceae + Dioscoreaceae]]

Pandanales

Velloziaceae Triuridaceae Stemonaceae [Pandanaceae + Cyclanthaceae]

Liliales

Corsiaceae [Campynemataceae [Petermanniaceae [Colchicaceae + Alstroemeriaceae]]

Melanthiaceae [[Philesiaceae + Rhipogonaceae] [Smilacaceae + Liliaceae]]]

Asparagales

Orchidaceae [[Boryaceae [Blandfordiaceae [Lanariaceae [Asteliaceae + Hypoxidaceae]]]]

[[Ixioliriaceae + Tecophilaeaceae] [Doryanthaceae [Iridaceae [Xeronemataceae

[Xanthorrhoeaceae [Alliaceae + Asparagaceae]]]]]]]

COMMELINIDS

Unplaced

Dasypogonaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

33

Arecales

Arecaceae

Poales

[Typhaceae + Bromeliaceae] [Rapateaceae [[Thurniaceae [Juncaceae + Cyperaceae]]

[[Mayacaceae [Eriocaulaceae + Xyridaceae]] [Flagellariaceae [Anarthriaceae

[Centrolepidaceae + Restionaceae]]] [Joinvilleaceae [Ecdeiocoleaceae + Poaceae]]]]]]]

Commelinales

[Commelinaceae + Hanguanaceae] [Philydraceae [Haemodoraceae + Pontederiaceae]]

Zingiberales

Musaceae [Strelitziaceae + Lowiaceae] Heliconiaceae [[Cannaceae + Marantaceae]

[Costaceae + Zingiberaceae]]

Ceratophyllales

Ceratophyllaceae

EUDICOTS

Ranunculales

Eupteleaceae [Papaveraceae [[[Lardizabalaceae + Circaeasteraceae] [Menispermaceae

[Berberidaceae + Ranunculaceae]]]]

Sabiales

Sabiaceae

Proteales

Nelumbonaceae [Platanaceae + Proteaceae]

Trochodendrales

Trochodendraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

34

Buxales

Haptanthaceae [Buxaceae + Didymelaceae]

CORE EUDICOTS

Gunnerales

Gunneraceae + Myrothamnaceae

Dilleniales

Dilleniaceae

Saxifragales

Peridiscaceae [[Paeoniaceae [Altingiaceae [Hamamelidaceae [Cercidiphyllaceae +

Daphniphyllaceae]]]] [[Crassulaceae [Aphanopetalaceae [Tetracarpaeaceae

[Penthoraceae + Haloragaceae]]]] [Iteaceae [Grossulariaceae + Saxifragaceae]]]]

Cynomoriaceae unplaced

Vitales

Vitaceae

ROSIDS

FABIDROSID I

Zygophyllales

Krameriaceae + Zygophyllaceae

Celastrales

Lepidobotryaceae + Celastraceae

Oxalidales

Huaceae [[Connaraceae + Oxalidaceae] [Cunoniaceae [Elaeocarpaceae [Brunelliaceae +

Cephalotaceae]]]]

Malpighiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

35

[Achariaceae [Goupiaceae [Violaceae + Passifloraceae] [Lacistemataceae Salicaceae]]

[Lophopyxidaceae Putranjivaceae] [Ctenolophonaceae [Erythroxylaceae +

Rhizophoraceae]] Linaceae Ixonanthaceae Humiriaceae Irvingiaceae

Centroplacaceae Pandaceae [[Bonnetiaceae + Clusiaceae] [Calophyllaceae

[Hypericaceae + Podostemaceae]]] [Malpighiaceae + Elatinaceae] Ochnaceae

[Peraceae [Rafflesiaceae + Euphorbiaceae]] [Phyllanthaceae + Picrodendraceae]

[Balanopaceae [[Trigoniaceae + Dichapetalaceae] [Chrysobalanaceae +

Euphroniaceae]]] Caryocaraceae

N-FIXING CLADE

Fabales

Quillajaceae [Fabaceae [Polygalaceae + Surianaceae]]

Rosales

Rosaceae [[Barbeyaceae [Elaeagnaceae Dirachmaceae Rhamnaceae]] [Ulmaceae

[Cannabaceae [Moraceae + Urticaceae]]]]

Cucurbitales

Anisophylleaceae [[Corynocarpaceae + Coriariaceae] [Cucurbitaceae [Tetramelaceae

[Datiscaceae + Begoniaceae]]]] Apodanthaceae

Fagales

Nothofagaceae [Fagaceae [[Myricaceae + Juglandaceae] [Casuarinaceae

[Ticodendraceae + Betulaceae]]]]

MALVIDROSID II

pgtGeraniales

Geraniaceae [[Melianthaceae + Francoaceae] [Vivianaceae + Ledocarpaceae]]

Myrtales

Combretaceae [[Onagraceae + Lythraceae] [[Vochysiaceae + Myrtaceae]

[Melastomataceae [Crypteroniaceae [Alzateaceae + Penaeaceae]]]]]

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

36

Crossosomatales

[Staphyleaceae [Guamatelaceae [Crossosomataceae + Stachyuraceae]]] [Aphloiaceae

[Geissolomataceae + Strasburgeriaceae]]

Picramniales

Picramniaceae

Sapindales

Biebersteiniaceae Nitrariaceae [Kirkiaceae [Anacardiaceae + Burseraceae]]

[Sapindaceae [Simaroubaceae Rutaceae Meliaceae]]

Huerteales

Gerradinaceae [Dipentodontaceae + Tapisciaceae]

Malvales

Neuradaceae [Thymelaeaceae [Sphaerosepalaceae Bixaceae [Cistaceae

[Sarcolaenaceae + Dipterocarpaceae]] [Cytinaceae + Muntingiaceae] Malvaceae]]

Brassicales

[Akaniaceae + Tropaeolaceae] [[Moringaceae + Caricaceae] [Setchellanthaceae

[Limnanthaceae [[Koeberliniaceae [Bataceae + Salvadoraceae]] [Emblingiaceae

[Pentadiplandraceae [Gyrostemonaceae Resedaceae Stixis and relatives] Tovariaceae

[Capparaceae [Cleomaceae + Brassicaceae]]]]]]]]

Berberidopsidales

Aextoxicaceae + Berberidopsidaceae

Santalales

Erythropalaceae Olacaceae [[Loranthaceae [Misodendraceae + Schoepfiaceae]]

[Opiliaceae + Santalaceae]] Balanophoraceae unplaced

Caryophyllales

[[Droseraceae [Nepenthaceae [Drosophyllaceae [Ancistrocladaceae +

Dioncophylleaceae]]]]] [[Frankeniaceae + Tamaricaceae] [Polygonaceae +

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

37

Plumbaginaceae]]] [Rhabdodendraceae [Simmondsiaceae [[Asteropeiaceae +

Physenaceae] [[Caryophyllaceae [Achatocarpaceae + Amaranthaceae]]

[Stegnospermataceae [Limeaceae [[Lophocarpaceae [Barbeuiaceae [Aizoaceae

[Sarcobataceae Phytolaccaceae Nyctaginaceae]]]] [Molluginaceae [Halophytaceae

Montiaceae Basellaceae Didiereaceae [Talinaceae [Portulacaceae [Anacampseros etc +

Cactaceae]]]]]]]]]]]]]

ASTERIDS

Cornales

[Cornaceae + Nyssaceae] [Hydrangeaceae + Loasaceae] Hydrostachyaceae [Curtisiaceae

+ Grubbiaceae]

Ericales

[Balsaminaceae [Marcgraviaceae + Tetrameristaceae]] [[Polemoniaceae +

Fouquieraceae] Lecythidaceae [[Sladeniaceae + Pentaphylacaceae] [Sapotaceae

[Ebenaceae [Maesaceae [Theophrastaceae [Myrsinaceae + Primulaceae]]]]]

[Mitrastemonaceae Theaceae [Symplocaceae [Styracaceae + Diapensiaceae]]

[[Sarraceniaceae [Roridulaceae + Actinidiaceae] [Clethraceae [Cyrillaceae +

Ericaceae]]]]]]

LAMIIDASTERID I

Unplaced

Metteniusaceae Oncothecaceae Icacinaceae and some unplaced ex-Icacinaceae

Garryales

Garryaceae + Eucommiaceae

Unplaced

Boraginaceae Vahliaceae

Gentianales

Rubiaceae [Gentianaceae [Loganiaceae [Gelsemiaceae + Apocynaceae]]]

Lamiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

38

Plocospermataceae [Oleaceae [Tetrachondraceae [Calceolariaceae

[Gesneriaceae Peltanthera [Pedaliaceae Carlemanniaceae Martyniaceae Stilbaceae

Plantaginaceae Linderniaceae Bignoniaceae Verbenaceae Lamiaceae Paulowniaceae

Schlegeliaceae Thomandersiaceae Phrymaceae Rehmannia Acanthaceae

Scrophulariaceae Orobanchaceae Byblidaceae Lentibulariaceae]]]]]

Solanales

[Montiniaceae [Sphenocleaceae + Hydroleaceae]] [Convolvulaceae + Solanaceae]

CAMPANULIDASTERID II

Aquifoliales

[[Cardiopteridaceae + Stemonuraceae] [Aquifoliaceae [Helwingiaceae +

Phyllonomaceae]]

Asterales

[Rousseaceae + Campanulaceae] Pentaphragmataceae [[Alseuosmiaceae [Phellinaceae

+ Argophyllaceae]] Stylidiaceae [Menyanthaceae [Goodeniaceae [Calyceraceae +

Asteraceae]]]]

Escalloniales

Escalloniaceae

Bruniales

[Bruniaceae + Columelliaceae]

Apiales

Pennantiaceae [Torricelliaceae [Griseliniaceae [Pittosporaceae [Araliaceae

[Myodocarpaceae + Apiaceae]]]]]

Paracryphiales

Paracryphiaceae

Dipsacales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

39

Adoxaceae [Diervillaceae [Caprifoliaceae [Linnaeaceae [Morinaceae [Dipsacaceae +

Valerianaceae]]]]]

LINKS TO ORDERS AND FAMILIES

[Back to Top]

MAIN GROUPINGS

Asterids commelinids core eudicots asterid 1 asterid

2 eudicots gymnosperms Magnoliophyta monocots N-fixing claderosids seed plants

ALPHABETICAL LISTING OF ALL ORDINAL NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | L | M | N | O | P | Q | R | S | T | U | V | W | X | Z

Abietales Acanthales Acerales Acorales Actinidiales Actinostrobales Adoxales Aescul

ales Agavales Aizoales AkanialesAlismatales Alliales Alseuosmiales Alstroemeriales

Altingiales Amaranthales Amaryllidales Amborellales AmbrosialesAmmiales Amomal

es Ancistrocladales Anisophylleales Annonales Anthobolales Apiales Apocynales Apo

nogetonalesAquifoliales Arales Araliales Aralidiales Araucariales Arecales Aristolochi

ales Athrotaxidales Asarales AsclepiadalesAsparagales Asphodelales Asparagales Ast

eliales Atriplicales Aucubales Austrobaileyales Avenales

Balanitales Balanopales Balanophorales Balsaminales Barbeyales Barclayales Batales

Begoniales BerberidalesBerberidopsidales Betulales Biebersteiniales Bignoniales Bix

ales Boraginales Brassicales Brexiales Bromeliales BrunialesBrunoniales Burmannial

es Burserales Butomales Buxales Byblidales

Cactales Callitrichales Calycanthales Calycerales Campanulales Campynematales Can

ellales Cannales CapparalesCaprifoliales Cardiopteridales Carduales Caricales Carle

manniales Caryophyllales Cassiales Casuarinales CelastralesCentrolepidales Cephalot

ales Cephalotaxales Ceratophyllales Cercidiphyllales Chenopodiales Chironiales Chlor

anthalesChrysobalanales Cinchonales Circaeasterales Cistales Citrales Cocosales Col

chicales Columelliales CombretalesCommelinales Connarales Convolvulales Coriarial

es Cornales Corylales Corynocarpales Crassulales CrossosomatalesCucurbitales Cunn

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

40

inghamiales Cunoniales Cupressales Cycadales Cyclanthales Cymodoceales Cynarales

CynomorialesCyperales Cyrillales Cytinales

Daphnales Daphniphyllales Dasypogonales Datiscales Degeneriales Desfontainiales D

iapensiales Didymelales DillenialesDioales Dioncophyllales Dioscoreales Diospyrales

Dipentodontales Dipsacales Droserales

Ebenales Echiales Elaeagnales Elaeocarpales Elatinales Elodeales Emmotales Empetr

ales Ephedrales Ericales EriocaulalesErythropalaless Escalloniales Eucommiales Euph

orbiales Eupomatiales Eupteleales Euryalaless

Fabales Fagales Falcatifoliales Ficales Flacourtiales Flagellariales Fouquieriales Franc

oales Frangulales

Galiales Garryales Geissolomatales Gentianales Geraniales Gesneriales Ginkgoales G

laucidiales Globulariales GnetalesGoodeniales Greyiales Griseliniales Grossulariales

Grubbiales Gunnerales Gyrocarpales Gyrostemonales

Haemodorales Haloragales Hamamelidales Hanguanales Heisteriales Helleborales He

lwingiales Himantandrales HippuridalesHomaliales Hortensiales Huales Huerteales

Hydatellales Hydnorales Hydrangeales Hydrastidales HydrocharitalesHydropeltidales

Hydrostachyales Hypericales Hypoxidales

Icacinales Illiciales Iridales Irvingales Iteales Ixiales

Jasminales Juglandales Julianiales Juncaginales Juncales

Lacistematales Lactoridales Lamiales Lardizabalales Laurales Lecythidales Ledocarpal

es Leitneriales LentibularialesLigustrales Liliales Limnanthales Linales Liriales Loasal

es Lobeliales Loganiales Lonicerales Loranthales LowialesLythrales

Magnoliales Malpighiales Malvales Marathrales Marcgraviales Mayacales Medusagy

nales Medusandrales MelanthialesMelastomatales Meliales Melianthales Meliosmal

es Menispermales Menyanthales Metteniusales MitrastemonalesMiyoshiales Moni

miales Moringales Musales Myricales Myristicales Myrothamnales Myrsinales Myrta

les

Najadales Nandinales Narcissales Nartheciales Nelumbonales Nepenthales Neuradal

es Nitrariales Nolanales NothofagalesNyctaginales Nymphaeales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

41

Ochnales Oenotherales Olacales Oleales Onagrales Oncothecales Opuntiales Orchid

ales Oxalidales

Saxifragales Pandales Pandanales Papaverales Paracryphiales Parasitaxales Paridales

Parnassiales PassifloralesPenaeales Pennantiales Pentaphragmatales Peridiscales P

etiveriales Petrosaviales Phellinales Philydrales PhyllanthalesPhysenales Phytolaccal

es Picramniales Pinales Pinguiculales Piperales Pittosporales Plantaginales Platanale

sPlumbaginales Poales Podocarpales Podophyllales Podostemales Polemoniales Pol

ygalales Polygonales PontederialesPortulacales Posidoniales Potamogetonales Primu

lales Proteales

Quercales Quillajales Quintiniales

Rafflesiales Ranunculales Rapateales Resedales Restionales Rhabdodendrales Rhamn

ales Rhinanthales RhizophoralesRhodorales Rhoipteleales Roridulales Rosales Rouss

eales Rubiales Ruppiales Rutales

Sabiales Salicales Salvadorales Samolales Samydales Sanguisorbales Santalales Sapin

dales Sapotales SarracenialesSaxegotheales Saxifragales Scheuchzeriales Sciadopitya

les Scleranthales Scrophulariales Scyphostegiales SedalesSilenales Simmondsiales S

milacales Solanales Sphenocleales Sphenostemonales Stangeriales Stellariales Stemo

nalesStilbales Stylidiales Styracales Surianales

Taccales Tamales Tamaricales Taxales Taxodiales Tecophilaeales Terebinthales Tern

stroemiales Theales TheligonalesThymelaeales Tiliales Tofieldiales Torricelliales Tov

ariales Tribelales Trilliales Trimeniales Triuridales TrochodendralesTropaeolales Turn

erales Typhales

Ulmales Urticales

Vacciniales Vahliales Vallisneriales Velloziales Veratrales Verbenales Viburnales Vinc

ales Violales Viscales VitalesVochysiales

Welwitschiales Winterales

Xanthorrhoeales Ximeniales Xyridales

Zamiales Zingiberales Zosterales Zygophyllales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

42

ALPHABETICAL LISTING OF ALL FAMILY NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | K | L | M | N | O | P | Q | R | S | T | U | V | W | X | Y

|Z

Back to Home

In some cases clicking on a family synonym will send you directly to the subfamily in

which the genus of the basionym is to be found

Abietaceae Abolbodaceae Abrophyllaceae Acalyphaceae Acanthaceae [Acanthochla

mydaceae] Acarnaceaee AceraceaeAchariaceae Achatocarpaceae Achradaceae Achy

ranthaceae Acmopylaceae Acoraceae Acristaceae ActaeaceaeActinidiaceae Actinota

ceae Amentotaxaceae Adenogrammaceae Adoxaceae Aegialitidaceae Aegicerataceae

Aegilopaceae Aeginetiaceaee Aegiphilaceae Aesculaceae Aextoxicaceae Agapanthac

eae Agavaceae AgdestidaceaeAgialidaceae Agrimoniaceae Agrostidaceae Ailanthace

ae Aitoniaceae Aizoaceae Akaniaceae Alangiaceae AlchemillaceaeAldrovandaceae A

lismataceae Alliaceae Allioniaceae Allophylaceae Aloaceae Alopecuraceae Alpiniacea

e AlseuosmiaceaeAlsinaceae Alsodeiaceae Alstroemeriaceae Altingiaceae Alzateacea

e Amaranthaceae Amaryllidaceae Amborellaceae Ambrosiaceae Amentotaxaceae A

mmanniaceae Ammiaceae Amomaceae Ampelidopsaceae Amygdalaceae Amyridacea

eAnacardiaceae Anagallidaceae Anarthriaceae Anchusaceae Ancistrocladaceae Andr

omedaceae AndropogonaceaeAndrostachyaceae Androsynaceae Anemarrhenaceae

Anemonaceae Angelicaceae Anisophylleaceae AnnonaceaeAnomochloaceae Anopter

aceae Anrederaceae Anthemidaceae Anthericaceae Anthobolaceae Antidesmataceae

Antirrhinaceae Antoniaceae Aparinaceae Aphanopetalaceae Aphloiaceae Aphyllanth

aceae Apiaceae ApocynaceaeApodanthaceae Aponogetonaceae Aporusaceae Apose

ridaceae Apostasiaceae Aptandraceae AquifoliaceaeAquilariaceae Aquilegiaceae A

raceae Aragoaceae Araliaceae Aralidiaceaee Araucariaceae Arbutaceae Arceuthidace

aeArceuthobiaceae Arctostaphylaceae Arctotidaceae Ardisiaceae Arecaceae Argoph

yllaceae Arisaraceae AristolochiaceaeAristoteliaceae Arjonaceae Armeriaceae Artem

isiaceae Arthrotaxidaceae Artocarpaceae Arundinaceae ArundinellaceaeAsaraceae A

sclepiadaceae Ascyraceaee Asiraceae Asparagaceae Asperulaceae Asphodelaceae As

pidistraceaeeAsteliaceae Asteraceae Asteranthaceae Asteropeiaceae Astragalaceae

Astrocarpaceae AthanasiaceaeAtherospermataceae Atriplicaceae Atropaceae Aucuba

ceae Aurantiaceae Austrobaileyaceae Austrotaxaceae AvenaceaeAverrhoaceae Avet

raceae Avicenniaceae Azaleaceae Azimaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

43

Balanitaceae Balanopaceae Balanophoraceae Balsameaceae Balsaminaceae Bambusa

ceae Barbaceniaceae BarbeuiaceaeBarbeyaceae Barclayaceae Barringtoniaceae Base

llaceae Bataceae Baueraceae Bauhiniaceae Baxteriaceae BegoniaceaeBehniaceae B

elangeraceae Belloniaceae Bembiciaceae Berberidaceae Berberidopsidaceae Berryac

eaee BersamaceaeBertyaceae Berzeliaceae Besleriaceae Betaceae Betulaceae Biebe

rsteiniaceae Bifariaceae Bignoniaceae BischofiaceaeBixaceae Blakeaceae Blandfordia

ceae Blattiaceae Blepharocaryaceae Blitaceae Blyxaceae Boerlagellaceae Bolivaracea

eBombacaceae Bonnetiaceae Bontiaceae Boopidaceae Boraginaceae Borassaceae B

oroniaceae BoryaceaeBotryodendraceae Bougainvilleaceae Boweniaceae Brassicacea

e Brachycaulaceae Bretschneideraceae BrexiaceaeBromeliaceae Brownlowiaceae Br

unelliaceae Bruniaceae Brunoniaceae Brunsvigiaceae Bryoniaceae BuchneraceaeBuci

daceae Buddlejaceae Buglossaceae Bulbocodiaceae Bumeliaceae Bupleuraceae Burc

hardiaceae BurmanniaceaeBurseraceae Butneriaceae Butomaceae Buxaceae Byblida

ceae Byttneriaceae

Cabombaceae Cacaoaceae Cactaceae Caesalpiniaceae Caladiaceae Calamaceae Calce

olariaceae CalectasiaceaeCalendulaceae Callaceae Callicomaceae Calligonaceae Calli

traceae Callitrichaceae Calochortaceae CalophyllaceaeCalthaceae Calycanthaceae Ca

lyceraceae Cambogiaceae Camelliaceae Campanulaceae CampynemataceaeCanacom

yricaceae Canellaceae Cannabaceae Cannaceae Canopodaceae Canotiaceae Cansjera

ceae CapparaceaeCaprariaceae Caprifoliaceae Cardiopteridaceae Carduaceae Carica

ceae Carissaceae Carlemanniaceae CarpinaceaeCarpodetaceae Cartonemataceae Ca

ryocaraceae Caryophyllaceae Casearieaceae Cassiaceae CassipoureaceaeCassythacea

e Castaneaceae Castelaceae Casuarinaceae Catesbaeaceae Cathedraceae Cecropiace

ae CedraceaeCedrelaceae Celastraceae Celosiaceae Celtidaceae Centaureaceae Cen

trolepidaceae Centroplacaceae CepaceaeCephalanthaceae Cephalotaceae Cephalota

xaceae Cerastiaceae Ceratoniaceae Ceratophyllaceae CerberaceaeCercidiphyllaceae

Cercocarpaceae Cercodiaceae Cereaceae Cerinthaceae Ceroxylaceae Cestraceae Cev

alliaceaeChailletiaceae Chamaedoraceae Chamaemoraceae Chamelauciaceae Chaun

ochitonaceae Chelidoniaceae ChelonaceaeChenopodiaceae Chimonanthaceae Chingi

thamnaceae Chionographidaceae Chiranthodendraceae ChironiaceaeChlaenaceae Ch

loanthaceae Chloranthaceae Chloridaceae Chlorogalaceae Chrysobalanaceae Cicerac

eae CichoriaceaeCimicifugaceae Cinchoniaceae Circaeaceae Circaeasteraceae Cissac

eae Cistaceae Citraceae Clematidaceae CleomaceaeClethraceae Clusiaceae Cneorac

eae Cnestidaceae Cnicaceae Cobaeaceae Cochlospermaceae Cocosaceae Coffeaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

44

Colchicaceae Coleogynaceae Columelliaceae Combretaceae Commelinaceae Comocl

adiaceae Compositae CompsoaceaeConantheraceae Connaraceae Conostylidaceae C

onvallariaceae Convolvulaceae Cordiaceae CoreopsidaceaeCoriandraceae Coriariacea

e Coridaceae Corispermaceae Cornaceae Corokiaceae Coronillaceae Correaceae Cor

rigiolaceaeCorsiaceae Corydalaceae Corylaceae Corynocarpaceae Coryphaceae Cost

aceae Cotyledonaceae CoulaceaeCoutariaceae Coutoubiaceae Crassulaceae Crescen

tiaceae Cressaceae Crinaceae Crocaceae CroomiaceaeCrossosomataceae Crotonacea

e Cruciferae Cryptaceae Crypteroniaceae Cryptocorynaceae CryptomeriaceaeCtenol

ophonaceae Cucurbitaceae Cunninghamiaceae Cunoniaceae Cupressaceae Curcumac

eae Curtisiaceae CuscutaceaeCyananthaceae Cyanastraceae Cynanchaceae Cyanella

ceae Cycadaceae Cyclanthaceae CyclantheraceaeCyclocheilaceae Cydoniaceae Cymo

doceaceae Cynaraceae Cynocrambaceae Cynomoriaceae Cyperaceae CyphiaceaeCyp

hocarpaceae Cypripediaceae Cyrillaceae Cyrtandraceae Cyrtanthaceae Cytinaceae

Dacrycarpaceae Dactylanthaceae Damasoniaceae Daphnaceae Daphniphyllaceae Das

ypogonaceae DatiscaceaeDaturaceae Daucaceae Davidiaceae Davidsoniaceae Decais

neaceae Deeringiaceae Degeneriaceae DelphiniaceaeDendrophthoaceae Columelliac

eae Detariaceae Dialypetalanthaceae Dianellaceae Dianthaceae DiapensiaceaeDicha

petalaceae Dichondraceae Diclidantheraceae Dicrastylidiaceae Dictamnaceae Didiere

aceae DidymelaceaeDidymocarpaceae Diegodendraceae Diervillaceae Digitalidaceae

e Dilatridaceae Dilleniaceae Dioaceae DionaeaceaeDioncophyllaceae Dioscoreaceae

Diosmaceae Diospyraceae Dipentodontaceae Diphylleiaceae Diplolaenaceae Dipsaca

ceaeDipterocarpaceae Dirachmaceae Disanthaceae Diselmaceae Dodonaeaceae Do

mbeyaceae Donatiaceae DorsteniaceaeDortmannaceae Doryanthaceae Drabaceae D

racaenaceae Dracontiaceae Drimyidaceae Droseraceae DrosophyllaceaeDryadaceae

Duabangaceae Duckeodendraceae Dulongiaceae Durantaceae Durionaceae Dysphani

aceae

Ebenaceae Eccremidaceae Ecdeiocoleaceae Echiaceae Echinopaceae Ehretiaceae Ela

eagnaceae ElaeocarpaceaeElegiaceae Elatinaceae Ellisiaceae Ellisiophyllaceae Elodea

ceae Elytranthaceae Embeliaceae Emblingiaceae EmmotaceaeEmpetraceae Encepha

lartaceae Engelhardtiaceae Enhalaceae Epacridaceae Ephedraceae Ephemeranthacea

eEpilobiaceae Epimediaceae Eragrostidaceae Eremolepidaceae Eremosynaceae Erica

ceae Erinaceae EriocaulaceaeEriogonaceae Eriospermaceae Erodiaceae Erycibaceae

Eryngiaceae Erysimaceae Erythroniaceae ErythropalaceaeErythrospermaceae Erythro

xylaceae Escalloniaceae Eschscholziaceae Eucomidaceae Eucommiaceae Eucryphiace

aeEuonymaceae Eupatoriaceae Euphorbiaceae Euphrasiaceae Euphroniaceae Eupo

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

45

matiaceae Eupteleaceae EuryalaceaeEustrephaceae Euthemidaceae Eutocaceae Exb

ucklandiaceae Exocarpaceae

Fabaceae Fagaceae Falcatifoliaceae Ferulaceae Festucaceae Ficaceae Ficoidaceae Fit

zroyaceae FlacourtiaceaeFlagellariaceae Flindersiaceae Foetidiaceae Forestieraceae

Fothergillaceae Fouquieriaceae Fragariaceae FrancoaceaeFrangulaceae Frankeniacea

e Fraxinaceae Fraxinellaceae Freycinetiaceae Fritillariaceae Fuchsiaceae Fumariacea

eFunkiaceae

Gaiadendraceae Galacaceae Galanthaceae Galaxiaceae Galedupaceae Galeniaceae G

aliaceae Garciniaceae GardeniaceaeGardneriaceae Garryaceae Geissolomataceae Ge

itonoplesiaceae Gelsemiaceae Geniostomaceae GentianaceaeGeonomataceae Geosir

idaceae Geraniaceae Gerrardinaceae Gesneriaceae Gethyllidaceae Gilliesiaceae Gina

lloaceaeGinkgoaceae Gisekiaceae Gladiolaceae Glaucidiaceae Glechomaceae Glinace

ae Globulariaceae Gnaphaliaceae GnetaceaeGoetzeaceae Gomortegaceae Gomphiac

eae Gomphrenaceae Gonystylaceae Goodeniaceae Gordoniaceae GouaniaceaeGoup

iaceae Gramineae Gratiolaceae Grewiaceae Greyiaceae Grielaceae Griseliniaceae Gr

onoviaceae GrossulariaceaeGrubbiaceae Guaiacanaceae Guamatelaceae Guettardac

eae Gunneraceae Gustaviaceae Guttiferae GyrocarpaceaeGyrostemonaceae

Hachetteaceae Haemanthaceae Haemodoraceae Halesiaceae Hallieraceae Halocarpa

ceae Halophilaceae HalophytaceaeHaloragaceae Hamamelidaceae Hanguanaceae H

aptanthaceae Harmandaceae Hebenstretiaceae HectorellaceaeHederaceae Hedyosm

aceae Hedyotidaceae Hedysaraceae Heisteriaceae Heleniaceae Heliamphoraceae H

elianthaceaeHelianthemaceae Helichrysaceae Heliconiaceae Helicteraceae Heliotropi

aceae Helleboraceae Heloniadaceae HelosidaceaeHelwingiaceae Hemerocallidaceae

Hemimeridaceae Hemiphylacaceae Henriqueziaceae Henslowiaceae Hermanniaceae

Hernandiaceae Herniariaceae Herreriaceae Hesperocallidaceae Heterantheraceae He

teropyxidaceae HeterostylaceaeHibbertiaceae Hibiscaceae Hilleriaceae Himantandra

ceae Hippocastanaceae Hippocrateaceae HippomanaceaeHippophaeaceae Hippurida

ceae Hirtellaceae Holacanthaceae Homaliaceae Hopkinsiaceae Hoplestigmataceae H

ordeaceaeHornschuchiaceae Hortensiaceae Hortoniaceae Hostaceae Houstoniaceae

Huaceae Huerteaceae HugoniaceaeHumbertiaceae Humiriaceae Hyacinthaceae Hyd

atellaceae Hydnoraceae Hydrangeaceae Hydrastidaceae HydrillaceaeHydrocharitacea

e Hydrocotylaceae Hydrogetonaceae Hydroleaceae Hydropeltidaceae Hydrophylacac

eae HydrophyllaceaeHydrostachyaceae Hymenocardiaceae Hypecoaceae Hyperanth

eraceae Hypericaceae Hypopityaceae HypoxidaceaeHypseocharitaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta

ceae ImperatoriaceaeInocarpaceae Inulaceae Ionidiaceae Iriarteaceae Iridaceae Irvi

ngiaceae Isnardiaceae Isophysidaceae IteaceaeStrasburgeriaceae Ixiaceae Ixioliriace

ae Ixonanthaceae

Jalapaceae Jamboliferaceae Japonoliriaceae Jasionaceae Jasminiaceae Johnsoniaceae

Joinvilleaceae JuglandaceaeJulianiaceae Juncaceae Juncaginaceae Juniperaceae Jus

siaeaceae Justiciaceae

Kadsuraceae Kaliphoraceae Kalmiaceae Kaniaceae Kiggelariaceae Kingdoniaceae Kin

giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae

Krameriaceae

Labiatae Lacandoniaceae Lachenaliaceae Lacistemataceae Lactoridaceae Lactucaceae

Lagerstroemiaceaee LamiaceaeLampsanaceae Lanariaceae Langsdorffiaceae Lantan

aceae Lapageriaceae Lardizabalaceae Lasiopetalaceae LauraceaeLawsoniaceae Laxm

anniaceae Lecythidaceae Ledaceae Ledocarpaceae Leeaceae Leguminosae Leitneriac

eae LemnaceaeLennoaceae Lentibulariaceae Lentiscaceae Leoniaceae Leonticaceae

Lepidobotryaceae LepidocarpaceaeLepidocaryaceae Lepidocerataceae Leptaulaceae

Leptospermaceae Lepturaceae Lepuropetalaceae LeuctenbergiaceaeLibocedraceae

Licaniaceae Ligustraceae Lilaeaceae Liliaceae Limeaceae Limnanthaceae Limnocharit

aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace

ae Linderniaceae Lindleyaceae Linnaeaceae LippayaceaeLiriaceae Liriodendraceae Li

ssocarpaceae Littorellaceae Loasaceae Lobeliaceaee Loganiaceae LomandraceaeLoni

ceraceae Lopeziaceae Lophiocarpaceae Lophiolaceae Lophiraceae Lophophytaceae L

ophopyxidaceae LoranthaceaeLotaceae Lowiaceae Lupulaceaee Luxemburgiaceae Lu

zuriagaceae Lychnidaceae Lyciaceae LyginiaceaeLygodisodeaceae Lysimachiaceae Ly

thraceae

Macarisiaceae Mackinlayaceae Madiaceae Maesaceae Magnoliaceae Malaceae Male

sherbiaceae MalortieaceaeMalpighiaceae Malvaceae Mangiaceae Manicariaceae Ma

paniaceae Marantaceae Marathraceae MarcgraviaceaeMartyniaceae Mastixiaceae

Matricariaceae Maundiaceae Mayacaceae Medeolaceae Medusagynaceae Peridiscac

eaeMelaleucaceae Melampyraceae Melanophyllaceae Melanthiaceae Melastomatac

eae Meliaceae Melianthaceae MelicaceaeMeliosmaceae Melittidaceae Melochiacea

e Memecylaceae Mendonciaceae Menispermaceae Menthaceae MenyanthaceaeMe

nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

47

Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

Nypaceae Nyssaceae

Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

ceae Papilionaceae Pappophoraceae PapyraceaeParacryphiaceae Parasitaxaceae Par

ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

48

hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

49

cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

50

ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 20: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

20

of the identification process before there can be further progress Instead those

characters that happen to be evident on a specimen can be used in whatever order is

convenient when linked to illustrations glossaries etc their power is enormous (see

Dallwitz et al 2000 [2006] for the principles underlying their construction and use)

Nevertheless dichotomous keys such as those of Hutchinson (1973) and Franz Thonner

(Geesink et al 1981) have their uses Of course taxa coming out adjacent in keys may

well not be at all related

(When identifying large numbers of plants even more efficient than either style of

identification and certainly lots more fun is sight identification Unless you have a

photographic memory you have to build up your knowledge of comparative plant

morphology - on which the ability to make accurate identifications depends - by

repeated observation When faced with an unknown plant I always look for leaf teeth

and stipules distinctive hair types especially stellate and T-shaped hairs glands and

punctations of any sort the presence of latex or other exudate and check leaf insertion

smelling crushed leaves can also be helpful In this context nodal anatomy can usually

be checked using a razor and a hand lens or even simply carefully examining leaf scars

The short paragraphs added after most families may help in confirming familial

identifications)

For the record and for the little that it is worth there are 4 orders and 13 families

of gymnosperms characterised on these pages and together they include some 82

genera and 947 species For angiosperms comparable figures are 56 orders 445

families 13208 genera and 261750 species (of which monocots include 11 orders 89

families 2759 genera and 52760 species) Note however that higher mathematics

was never my strong point and anyway these are pretty meaningless figures even for

species which many (but not all) might concede smacked slightly more of reality than

other taxa estimates range as high as 422000 (Govaerts 2001) Furthermore numbers

of genera and species change daily Nevertheless as emphasized here families are

useful in teaching we as a community can ensure that their limits remain largely stable

and by concentrating on relatively few of them one can gain some familiarity with much

of the worlds flora For summaries of the sizes of orders and families and of the general

arrangement followed in these pages see also the Statistics page itself

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

21

A PHYLOGENETIC TREE OF SEED PLANTS (THE MAIN TREE)

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

22

The Main Tree above is a conservative summary of well supported relationships

between the major clades of extant seed plants Reading - interpreting and

understanding - such a phylogenetic tree is not necessarily easy so reading a tree may

require a bit of practice OHara (eg 1988 1992) provides an accessible introduction to

thinking and talking about history in the context of looking at phylogenetic trees More

recently Krell and Cranston (2004) Crisp and Cook (2005) and others have emphasized

how careful one must be when interpreting and talking about evolution in ladderized

trees in particular and phylogenetic trees in general Gregory (2008) provides a

particularly detailed discussion of this problem In ladderized trees the smaller (in terms

of numbers of terminals) sister taxon is consistently shown on the same side at every

node thus the trees tend to be pectinate like the teeth of a comb however the

horizontal axis that is as a result so evident has no polarity or particular direction The

use of the adjective basal when discussing phylogenetic trees is especially dangerous

(see also Wojciechowski et al 2004 D Soltis et al 2005b) When I use the term and the

context is not otherwise clear I am referring to the pectinations at the base of a

ladderised tree Remember that when talking about sister taxa one can never be basal

to the other or older or younger although members of one can be more derived - in the

sense of having more apomorphies - than the other (but even then one has to be

careful) The words primitive and advanced and lower and higher should

practically never be used especially when talking about taxa Amborellaceae and

Pinaceae are sister to all other flowering plants and Pinales and are basal to the crown

groups of all other flowering plantsPinales respectively but that does not mean they

are thereby primitive Pinaceae in particular have numerous apomorphies The word

plesiomorphic is far less loaded than primitive and can be used to talk about

individual characters

Most trees here have been more or less ladderized as is the one above The

ladderization may be imperfect for example you can see that on the Main Tree

asterids with ten orders follow rosids and relatives which have seventeen although

this is in part because of the currently unresolved nature of relationships between the

main clades of core eudicots (this is likely to change soon) The Main Tree could have

been drawn with Amborella Acorus or a host of other taxa at the far right without

offending any relationships phylogenetic trees are like mobiles the only fixed points

being the nodes Despite the problems just mentioned pectination interpreted

carefully has its value and it is not an arbitrary process As one reads the terminals of a

pectinate tree from left to right adjacent terminals will be separated by apomorphies

that will cumulate in an appropriate fashion When thinking of a book or a herbarium

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

23

sequence (see Haston et al 2007) this is of some value Nymphaeales and

Austrobaileyales are here adjacent on the tree but they could be separated by

hundreds of families in the sequence merely by rotating only the node from which

Nymphaeales and their sister group (all other angiosperms minus about 75 species)

arise If adjacent in a book or herbarium or as in these summary pages then it is

relatively easy to relate their characters both apomorphic and plesiomorphic but if

separated by hundreds of pages or two floors in a large building then it is less easy to

get anything from the sequence (Since all orders in the pages of the main part of this

site are preceded by the apomorphies of all nodes immediately below them in the seed

plant phylogeny and because of the linkages that have been built in to the site this

problem is much less serious there there is no sequence) So the particular sequence of

all seed plants that is based on a phylogenetic classification but is to be used in herbaria

or any other place where taxa have to be arranged linearly as in this part of APweb

will be one that maximizes the number of taxa that are both successive branches of the

tree and placed successively in the sequence Since specimens are generally filed under

families the outline of a new family sequence for arranging herbaria and books can now

be suggested (Haston et al 2007) and such a standardized sequence will further help

teaching and learning about plants

ON THINKING ABOUT APOMORPHIES

Identifying apomorphies is important because understanding the evolution of

morphology in the broad sense - ie including anatomy chemistry etc - in the context

of diversification is one of our major goals For identifying apomorphies several

preconditions must be met One needs to have an accurate well supported phylogeny

one has to have examined the right taxa both from the point of view of morphology and

molecules one has to have coded the characters right (ie delimited states

appropriately) and one has to use an appropriate model of evolution when fitting the

variation to the tree (see eg Omland 1999 Stevens 2006b) If the distributions of

apomorphies below are compared with those in other studies (eg Turgeon et al 2001

Bremer et al 2001 Endress 2001 Albach et al 2001a Judd amp Olmstead 2004 D Soltis

et al 2005b Zhang et al 2006 Judd et al 2007) the reader may find a number of

differences Although I have integrated such studies as far as is possible there are five

reasons that link to the preconditions just mentioned why there may be differences

where features are placed on trees

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

24

1 I may not have found all the information about a particular character there may be

disagreement over its interpretation or I have added information

2 The sampling of nearly all molecular studies is very incomplete (see Salisbury amp Kim

2001 for problems caused by sampling) indeed it is commonplace to decry the

incompleteness of molecular sampling But not only is the sampling in molecular studies

often less than we might wish that of the morphological and chemical characters whose

evolution we are interested in understanding is also often very poor So for many

anatomical chemical and embryological characters that are confidently said to

characterise families and other groups we all too often have no idea if those characters

are applicable to the whole clade or just to a subgroup within that clade However as

our sampling improves we can locate changes on the tree more precisely if in

unexpected positions Thus is is sometimes suggested that Ericaceae have ellagic acid

(Soltis et al 2005b) in fact the little evidence we have indicates that only a small clade

of some 80 species has ellagic acid and the rest of the family - itself alone about one

third of all Ericales - does not Similarly Ericaceae are noted for their

ectendomycorrhizal associations but Enkianthus sister to all other Ericaceae

apparently lacks these (Abe 2005) Thus these mycorrhizae and their associated

distinctive hair roots are unlikely to be apomorphies of Ericaceae - however it appears

that other families near Ericaceae may also have ectendomycorrhizae (see Asai 1934)

3 I am fitting characters to a very conservative tree with many polytomies although the

nodes that are utilised are for the most part strongly supported polytomies make the

optimisation of characters that is the assigment of character state change to a

particular node on the tree notably difficult (eg Madison amp Madison 2002) In nearly

all studies of the evolution of characters distributions of characters are optimised on a

more or less fully resolved tree and the construction of supertrees may yield yet more

detailed hypotheses of relationships (for literature on supertrees see Cotton amp

Wilkinson 2007 2008) Of course some nodes on such fully resolved trees andor

supertrees may have little support and optimisations of characters on such trees may

carry correspondingly little conviction Even parts of some of the trees used here have

poor support eg relationships within aquatic Alismatales etc although I indicate

when this is the case

4 Exactly how one goes about optimising a character on a tree is critically important

Even using simple parsimony optimisations (ACCTRAN or DELTRAN ACCelerated

TRANsitions or DELayed TRANsitions) the position of synapomorphies on trees - and

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

25

hence our ideas of evolution - may differ and this is still more true if one uses maximum

likelihood or Bayesian methods and makes apparently reasonable suggestions about

weighting gains over losses (eg Donoghue amp Ackerley 1996 Cunningham et al 1998

Omland 1997 1999 Ree amp Donoghue 1999 Polly 2001 Webster amp Purvis 2001

Ronquist 2004 Crisp amp Cook 2005) Sannier et al (2007) show how in in Arecaceae that

where on a phylogenetic tree one might peg changes in microsporogenesis will depend

on the methods one uses to do this and Pedersen et al (2007) discuss the sometimes

very substantial effect of node support on the posterior probabilities of ancestral

character states Here I use parsimony optimization not always as explicit as it might be

but I have often indicated where there are particularly important uncertainties as to the

positions of particular character changes on the tree

5 Finally although I have paid quite a lot of attention to the delimitation of the

character states that make up all the characterizations I have not spent enough time on

this critical operation If we are interested in understanding evolution then fitting the

basic variation - not character states - to a tree in principle allows greater flexibility in

understanding morphology in the context of local phylogenies (see also Stevens 2000

Endress 2005c) However many character states used here are delimited globally that

is they are circumscribed in the context of the variation shown by individual characters

across all angiosperms andor in the context of classic ideas of character evolution

Character states often have arbitrary limits and serve best to communicate

information whether they are in fact suitable for either phylogenetic analysis or

understanding evolution are separate issues Studies have rather unsurprisingly

perhaps but importantly shown that dividing the one character into different sets of

states may yield differing ideas of evolution of that character (eg Lamb Frye amp Kron

2003 Hibbett 2004) When looking at trees on which character states are optimised

one should bear in mind the problems surrounding the delimitation of states (eg

Stevens 2000 2006b) and the danger of using pollen or other types - constructs

based on many characters that vary independently but which effectively get lost in

these types thus Blackmore et al (2009) decomposed the pollen types in Asteraceae

into 52 characters

I use Remanes three main criteria of homology or better similarity when

determining the basic similarity of structures on different organisms (see Remane 1952)

These criteria are special properties position and intermediates Special properties

include anatomical or chemical characters gene expression data etc Position refers

to the position of an organ with respect to landmarks on the plant Although plants are

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

26

plants and landmarks are not as fixed as one might like trying to work out the relative

positions of parts is a good way of understanding morphology so long as one does not

try to out-do Procrustes Intermediates include intermediates found on the same

plant or on different plants Intermediates may observed during development or be

adult structures Thus one may find changes during development which suggest what

the nature of a particular structure is and very different stuctures on different plants

may turn out to be similar early in development Or one can simply compare different

taxa and see that structures that appear to be very different when comparing two

different taxa can be linked morphologically by looking at other taxa

Nevertheless the use of these criteria may not yield an unambiguous answer as to

what a structure is even given a solid phylogeny and an improved understanding of

development (see Jaramillo amp Kramer 2007 for a useful discussion) As Endress (2005c)

observed a number of features - position function development shape anatomy

histology gene activity and relationships to other taxa that clearly have petals - can be

used to distinguish a petal (for example) from other floral structures if a petal does not

have one of these features is it thereby not a petal Thus Maturen et al (2005) recently

found that floral organ diversity genes (B and C) were expressed in the large white

inflorescence bracts of Cornus (see also Costa et al 2005) Peney et al (2005) noted that

not all monosulcate pollen grains in monocots have the same developmental pathway

and that as a result such pollen might not have the same ancestral state Reeves and

Olmstead (2003) suggested that the genetic mechanisms causing monosymmetry in

Lamiales and Solanales were different and Serna and Martin (2006) described similar

problems with the development of hairs in Arabidopsis when compared with that of

hairs in Antirrhinum and Solanaceae Indeed as one perhaps might expect delimitation

of states and characters does not necessarily become easier with increasing knowledge

of development etc Thus Buzgo et al (2004) Matthews and Endress (2005) and others

have shown how hard it can be to distinguish between eg prophylls and other floral

structures as their behaviour is studied during the course of the development of the

flower Similarly at what concentration is a particular secondary metabolite deemed to

be present (Waterman (2007)

The validity of the approach used here that of fitting morphological variation to a

largely molecular-based tree may be questioned However I think it rather unlikely that

well-supported molecular branches will be overturned by morphological data Indeed

analyses of morphological data alone do provide support for many of the clades evident

in molecule-only analyses and analyses with morphological and molecular data

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

27

together may lead to increased support for clades (eg Hufford 1992 Nandi et al 1998

[but adding morphological data reduces support for a number of critical clades too]

Doyle amp Endress 2000 however in none of these papers is the use of morphology

without ambiguity) It is unfortunately clear that the use of morphology alone may not

suggest problems in the phylogenetic placement of taxa that later turn out to have been

wrongly included (eg Zhang et al 1992) Thus I have been wary of putting much weight

on clades that have only morphological support but note that I have treated molecule-

based clades with low bootstrap or jacknife support values (esp below 70) or low

posterior probabilies (below 095) likewise Although I may have been mistaken in

placing so much emphasis on molecular data in terms of providing the basic

phylogenetic framework for angiosperms morphological and molecular data are only

very rarely in irreconcilably strong conflict There are indeed a few places where the

conflict seems extreme These include the relative positions of the Monimiaceae and

Hernandiaceae (Laurales) the position of Hanguanaceae (Commelinales [as here] or

Zingiberales) and of Triplostegia (is it in Dipsacaceae or Valerianaceae - see

Dipsacales) Fossils are unlikely to affect the topologies of the trees presented here but

see below for their importance in understanding morphological evolution in general and

the evolution of angiosperms in particular Nevertheless some largely reject the idea

that trees based on molecular data alone can recover phylogenetic relationships

especially when branching points are old and prefer to used trees based on analysis of

morphological data including those taken from fossils (Hilton amp Bateman 2006 Farjon

2007)

All in all however the extent of the congruence between morphological and

molecular data is impressive and heartening and many clades can be characterised

morphologically It seemed in 1998 that there were no unambiguous morphological

synapomorphies for angiosperm orders (K Bremer 2000) and this is still true if by

unambiguous is meant non-homoplasious However many orders and other clades

have synapomorphies even if these may be indistinguishable at least at the current

level of morphological and developmental knowledge from parallel occurences

elsewhere As our morphological knowledge increases so too does the number of

apomorphies

Endress and Matthews (2006a) emphasize the importance of tendencies and

developmental constraints when thinking about characters of clades within the rosids -

many characters or character combinations occur in a rather sporadic fashion within

the clade and apparently notably less frequently outside it There are a number of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

28

examples of tendencies here and in the individual order pages such as the features

enclosed by parentheses in groups above the level of family eg see (cuticular waxes

as aggregated rodlets) for the commelinids and the discussion of the distribution of

polyandry that is flowers with numerous stamens in the asterid I + II groups etc

Tendencies may involve more than single characters When thinking of the

associations of features like integument thickness and vascularization of the integument

(Endress amp Matthews 2006a) one can imagine fairly simple developmental

preconditions being operative As Endress and Matthews (2006a) note it is difficult to

imagine vascular tissue developing in an integument only two cells thick Changes here

would be loosely correlated if morphologically linked However the strongly correlated

changes noted by Givnish et al (2005) are ecologically linked but are presumably

morphologicallydevelopmentally independent When there seem to be characters

evolving more or less together tests can be carried out to see if the changes are

concentrated on certain branches of the tree (eg Maddison 1990 Sanderson 1991

Maddison amp Maddison 2000) Such tests have rarely been carried out at higher levels in

angiosperms

One of the most striking examples of a tendency is the distribution of N-fixation

restricted as it is to a monophyletic group of four clades although it has arisen seven (or

perhaps several more) times independently within the N-fixing clade and several

members of two quite different kinds of bacteria are involved (eg Clawson et al 2004

Elliott et al 2007 Sprent amp James 2007) Other examples of work that bears on the issue

of tendencies include the findings that flowers of polysymmetrical Arabidopsis have

genes like TCP1 that are expressed asymmetrically during early development and TCP1

is a probable orthologue of the well-known CYC gene of Antirrhinum that is involved in

the development of monosymmetric slowers there (Cubas et al 2001 Costa et al 2005

etc) Parallelism might build on this underlying morphologically cryptic monosymmetry

even if details of the genetic mechanisms causing the monosymmetry evident in

particular groups may be different (Reeves amp Olmstead 2003 Cubas 2004 see above)

Similarly there have been several recent suggestions that the capability to synthesise a

particular metabolite may be switched off but not lost and so can sometimes be

reacquired (eg Wink amp Witte 1983 Wink 2003 Liscombe et al 2005 Larsson 2007

Waterman 2007) Hence perhaps the rather spotty distribution of many secondary

metabolites like ellagic acid the indole alkaloid camptothecin iridoids etc when

considered in the context of phylogenies As a non-botanical example - but a rather nice

one - Salwini-Plawen and Mayr (1961) suggested some time ago that there has been

considerable parallelism (40-65 or more independent origins) in the evolution of eyes in

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

29

metazoans However Pax 6 seems to be a master control gene that is involved in eye

formation perhaps throughout all bilateralians (eg Gehring amp Ikeo 1999 Erwin amp

Davidson 2002) which changes the nature of the problem Note however that in

general our invocation of evolutionary tendencies to explain the patterns of

distributions of characters is really an admission that we do not understand those

patterns

As we find out more about variation we will probably find fewer and fewer features

constant throughout a group Most unqualified statements of presence and absence

should properly be qualified as usually present or usually absent if one is thinking of

the characterisations on the order pages as encompassing the total variation within a

clade Thus Pistia alone among monocots as so far known has sieve tube plastids with

starch grains not protein crystals However this does not affect the fact that sieve tube

plastids with cuneate protein crystals and lacking starch grains are an apomorphy for

monocots Furthermore fossils in a number of cases suggest character combinations

unknown in extant taxa as may be seen in the discussions of Fagaceae Platanaceae

Iteaceae Calycanthaceae etc Confusing the issue there may be questions as to where

exactly on the tree a particular fossil is to be placed (eg see Nymphaeaceae

Calycanthaceae Archaefructus etc)

To summarize given our current understandings of both phylogenies and

characters evolution of some characters in which we are interested seems very labile

(see eg D Soltis et al 2005b Endress and Matthews 2006a Stevens 2006b Ekman et

al 2008 for a good recent study) and I have been cautious when talking about character

evolution Much effort must continue to be spent in summarizing characters of well-

established clades at all levels providing features by which they may be recognized and

signaling synapomorphies Remember that (1) the basic morphological anatomical and

chemical knowledge of many critical taxa is woefully incomplete (2) different

assumptions about character evolution may greatly affect the position of

synapomorphies on trees (3) in many cases relationships within and between many

groups are too uncertain at present to worry very much about synapomorphies and

(4) we must be clear about what we do and do not not know As mentioned above I

have indicated in a number of places where there are particularly important

uncertainties as to where characters should be placed on the tree Nevertheless it is a

relatively easy matter to update notes such as these and it can be a simple matter to

incorporate new data on characters that have never before been considered in the

context of a tree Much basic - and unfortunately perhaps unfashionable - work must be

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

30

carried to clarify the distribution of morphological anatomical and chemical characters

for examples of what can to be done see the work of P K Endress and collaborators

(floral morphology and development) and S R Jensen and collaborators (iridoids) But

acquiring information about nodal anatomy cell and tissue distribution nodal anatomy

and the like is not difficult at all and should be generally encouraged

But all this is merely a necesary prelude to the understanding of evolution For this

one needs to know a lot more including dating the phylogeny understanding the

function(s) of characters factoring in relevant aspects of the palaeoenvironment in

which an apomorphy first appeared etc

SUMMARY OF THE SYSTEM

Below is a formalised summary of the relationships within orders of the families of seed

plants There are a few families that are not recognised even as options in APG II and

vice versa as well as a few extra orders The families to be recognised in the most

recent edition of Mabberleys The Plant Book (Mabberley 2008) are also largely

consistent with those below But all differences are trivial and will - I hope - eventually

disappear

Square brackets - [] - enclose clades the plus sign - + - designates sister taxa a

comma - - denotes part of a polytomy and quotation marks - - denotes a

paraphyletic group

SEED PLANTS

GYMNOSPERMS

Cycadales

Cycadaceae + Zamiaceae

Ginkgoales

Ginkgoaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

31

Pinales

Pinaceae [[Araucariaceae [Podocarpaceae + Phyllocladaceae]] [Sciadopityaceae

[Taxaceae + Cupressaceae]]]

Gnetales

Ephedraceae [Gnetaceae + Welwitschiaceae]

ANGIOSPERMSFLOWERING PLANTS

Amborellales

Amborellaceae

Nymphaeales

Hydatellaceae [Cabombaceae + Nymphaeaceae]

Austrobaileyales

Austrobaileyaceae [Schisandraceae + Trimeniaceae]

Chloranthales

Chloranthaceae

MAGNOLIIDS

Magnoliales

Myristicaceae [Magnoliaceae [[Himantandraceae + Degeneriaceae] [Eupomatiaceae +

Annonaceae]]]

Laurales

Calycanthaceae [[Siparunaceae [Gomortegaceae + Atherospermataceae]] [Monimiaceae

[Hernandiaceae + Lauraceae]]]

Canellales

Canellaceae + Winteraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

32

Piperales

[Hydnoraceae + Aristolochiaceae] [Piperaceae + Saururaceae]

MONOCOTS

Acorales

Acoraceae

Alismatales

Araceae [Tofieldiaceae [[Alismataceae [Hydrocharitaceae + Butomaceae]]

[Scheuchzeriaceae [Aponogetonaceae [Juncaginaceae [Maundiaceae [[Posidoniaceae

[Ruppiaceae + Cymodoceaceae]] [Zosteraceae + Potamogetonaceae]]]]]]]

Petrosaviales

Petrosaviaceae

Dioscoreales

Nartheciaceae [[Taccaceae + Thismiaceae] [Burmanniaceae + Dioscoreaceae]]

Pandanales

Velloziaceae Triuridaceae Stemonaceae [Pandanaceae + Cyclanthaceae]

Liliales

Corsiaceae [Campynemataceae [Petermanniaceae [Colchicaceae + Alstroemeriaceae]]

Melanthiaceae [[Philesiaceae + Rhipogonaceae] [Smilacaceae + Liliaceae]]]

Asparagales

Orchidaceae [[Boryaceae [Blandfordiaceae [Lanariaceae [Asteliaceae + Hypoxidaceae]]]]

[[Ixioliriaceae + Tecophilaeaceae] [Doryanthaceae [Iridaceae [Xeronemataceae

[Xanthorrhoeaceae [Alliaceae + Asparagaceae]]]]]]]

COMMELINIDS

Unplaced

Dasypogonaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

33

Arecales

Arecaceae

Poales

[Typhaceae + Bromeliaceae] [Rapateaceae [[Thurniaceae [Juncaceae + Cyperaceae]]

[[Mayacaceae [Eriocaulaceae + Xyridaceae]] [Flagellariaceae [Anarthriaceae

[Centrolepidaceae + Restionaceae]]] [Joinvilleaceae [Ecdeiocoleaceae + Poaceae]]]]]]]

Commelinales

[Commelinaceae + Hanguanaceae] [Philydraceae [Haemodoraceae + Pontederiaceae]]

Zingiberales

Musaceae [Strelitziaceae + Lowiaceae] Heliconiaceae [[Cannaceae + Marantaceae]

[Costaceae + Zingiberaceae]]

Ceratophyllales

Ceratophyllaceae

EUDICOTS

Ranunculales

Eupteleaceae [Papaveraceae [[[Lardizabalaceae + Circaeasteraceae] [Menispermaceae

[Berberidaceae + Ranunculaceae]]]]

Sabiales

Sabiaceae

Proteales

Nelumbonaceae [Platanaceae + Proteaceae]

Trochodendrales

Trochodendraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

34

Buxales

Haptanthaceae [Buxaceae + Didymelaceae]

CORE EUDICOTS

Gunnerales

Gunneraceae + Myrothamnaceae

Dilleniales

Dilleniaceae

Saxifragales

Peridiscaceae [[Paeoniaceae [Altingiaceae [Hamamelidaceae [Cercidiphyllaceae +

Daphniphyllaceae]]]] [[Crassulaceae [Aphanopetalaceae [Tetracarpaeaceae

[Penthoraceae + Haloragaceae]]]] [Iteaceae [Grossulariaceae + Saxifragaceae]]]]

Cynomoriaceae unplaced

Vitales

Vitaceae

ROSIDS

FABIDROSID I

Zygophyllales

Krameriaceae + Zygophyllaceae

Celastrales

Lepidobotryaceae + Celastraceae

Oxalidales

Huaceae [[Connaraceae + Oxalidaceae] [Cunoniaceae [Elaeocarpaceae [Brunelliaceae +

Cephalotaceae]]]]

Malpighiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

35

[Achariaceae [Goupiaceae [Violaceae + Passifloraceae] [Lacistemataceae Salicaceae]]

[Lophopyxidaceae Putranjivaceae] [Ctenolophonaceae [Erythroxylaceae +

Rhizophoraceae]] Linaceae Ixonanthaceae Humiriaceae Irvingiaceae

Centroplacaceae Pandaceae [[Bonnetiaceae + Clusiaceae] [Calophyllaceae

[Hypericaceae + Podostemaceae]]] [Malpighiaceae + Elatinaceae] Ochnaceae

[Peraceae [Rafflesiaceae + Euphorbiaceae]] [Phyllanthaceae + Picrodendraceae]

[Balanopaceae [[Trigoniaceae + Dichapetalaceae] [Chrysobalanaceae +

Euphroniaceae]]] Caryocaraceae

N-FIXING CLADE

Fabales

Quillajaceae [Fabaceae [Polygalaceae + Surianaceae]]

Rosales

Rosaceae [[Barbeyaceae [Elaeagnaceae Dirachmaceae Rhamnaceae]] [Ulmaceae

[Cannabaceae [Moraceae + Urticaceae]]]]

Cucurbitales

Anisophylleaceae [[Corynocarpaceae + Coriariaceae] [Cucurbitaceae [Tetramelaceae

[Datiscaceae + Begoniaceae]]]] Apodanthaceae

Fagales

Nothofagaceae [Fagaceae [[Myricaceae + Juglandaceae] [Casuarinaceae

[Ticodendraceae + Betulaceae]]]]

MALVIDROSID II

pgtGeraniales

Geraniaceae [[Melianthaceae + Francoaceae] [Vivianaceae + Ledocarpaceae]]

Myrtales

Combretaceae [[Onagraceae + Lythraceae] [[Vochysiaceae + Myrtaceae]

[Melastomataceae [Crypteroniaceae [Alzateaceae + Penaeaceae]]]]]

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

36

Crossosomatales

[Staphyleaceae [Guamatelaceae [Crossosomataceae + Stachyuraceae]]] [Aphloiaceae

[Geissolomataceae + Strasburgeriaceae]]

Picramniales

Picramniaceae

Sapindales

Biebersteiniaceae Nitrariaceae [Kirkiaceae [Anacardiaceae + Burseraceae]]

[Sapindaceae [Simaroubaceae Rutaceae Meliaceae]]

Huerteales

Gerradinaceae [Dipentodontaceae + Tapisciaceae]

Malvales

Neuradaceae [Thymelaeaceae [Sphaerosepalaceae Bixaceae [Cistaceae

[Sarcolaenaceae + Dipterocarpaceae]] [Cytinaceae + Muntingiaceae] Malvaceae]]

Brassicales

[Akaniaceae + Tropaeolaceae] [[Moringaceae + Caricaceae] [Setchellanthaceae

[Limnanthaceae [[Koeberliniaceae [Bataceae + Salvadoraceae]] [Emblingiaceae

[Pentadiplandraceae [Gyrostemonaceae Resedaceae Stixis and relatives] Tovariaceae

[Capparaceae [Cleomaceae + Brassicaceae]]]]]]]]

Berberidopsidales

Aextoxicaceae + Berberidopsidaceae

Santalales

Erythropalaceae Olacaceae [[Loranthaceae [Misodendraceae + Schoepfiaceae]]

[Opiliaceae + Santalaceae]] Balanophoraceae unplaced

Caryophyllales

[[Droseraceae [Nepenthaceae [Drosophyllaceae [Ancistrocladaceae +

Dioncophylleaceae]]]]] [[Frankeniaceae + Tamaricaceae] [Polygonaceae +

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

37

Plumbaginaceae]]] [Rhabdodendraceae [Simmondsiaceae [[Asteropeiaceae +

Physenaceae] [[Caryophyllaceae [Achatocarpaceae + Amaranthaceae]]

[Stegnospermataceae [Limeaceae [[Lophocarpaceae [Barbeuiaceae [Aizoaceae

[Sarcobataceae Phytolaccaceae Nyctaginaceae]]]] [Molluginaceae [Halophytaceae

Montiaceae Basellaceae Didiereaceae [Talinaceae [Portulacaceae [Anacampseros etc +

Cactaceae]]]]]]]]]]]]]

ASTERIDS

Cornales

[Cornaceae + Nyssaceae] [Hydrangeaceae + Loasaceae] Hydrostachyaceae [Curtisiaceae

+ Grubbiaceae]

Ericales

[Balsaminaceae [Marcgraviaceae + Tetrameristaceae]] [[Polemoniaceae +

Fouquieraceae] Lecythidaceae [[Sladeniaceae + Pentaphylacaceae] [Sapotaceae

[Ebenaceae [Maesaceae [Theophrastaceae [Myrsinaceae + Primulaceae]]]]]

[Mitrastemonaceae Theaceae [Symplocaceae [Styracaceae + Diapensiaceae]]

[[Sarraceniaceae [Roridulaceae + Actinidiaceae] [Clethraceae [Cyrillaceae +

Ericaceae]]]]]]

LAMIIDASTERID I

Unplaced

Metteniusaceae Oncothecaceae Icacinaceae and some unplaced ex-Icacinaceae

Garryales

Garryaceae + Eucommiaceae

Unplaced

Boraginaceae Vahliaceae

Gentianales

Rubiaceae [Gentianaceae [Loganiaceae [Gelsemiaceae + Apocynaceae]]]

Lamiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

38

Plocospermataceae [Oleaceae [Tetrachondraceae [Calceolariaceae

[Gesneriaceae Peltanthera [Pedaliaceae Carlemanniaceae Martyniaceae Stilbaceae

Plantaginaceae Linderniaceae Bignoniaceae Verbenaceae Lamiaceae Paulowniaceae

Schlegeliaceae Thomandersiaceae Phrymaceae Rehmannia Acanthaceae

Scrophulariaceae Orobanchaceae Byblidaceae Lentibulariaceae]]]]]

Solanales

[Montiniaceae [Sphenocleaceae + Hydroleaceae]] [Convolvulaceae + Solanaceae]

CAMPANULIDASTERID II

Aquifoliales

[[Cardiopteridaceae + Stemonuraceae] [Aquifoliaceae [Helwingiaceae +

Phyllonomaceae]]

Asterales

[Rousseaceae + Campanulaceae] Pentaphragmataceae [[Alseuosmiaceae [Phellinaceae

+ Argophyllaceae]] Stylidiaceae [Menyanthaceae [Goodeniaceae [Calyceraceae +

Asteraceae]]]]

Escalloniales

Escalloniaceae

Bruniales

[Bruniaceae + Columelliaceae]

Apiales

Pennantiaceae [Torricelliaceae [Griseliniaceae [Pittosporaceae [Araliaceae

[Myodocarpaceae + Apiaceae]]]]]

Paracryphiales

Paracryphiaceae

Dipsacales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

39

Adoxaceae [Diervillaceae [Caprifoliaceae [Linnaeaceae [Morinaceae [Dipsacaceae +

Valerianaceae]]]]]

LINKS TO ORDERS AND FAMILIES

[Back to Top]

MAIN GROUPINGS

Asterids commelinids core eudicots asterid 1 asterid

2 eudicots gymnosperms Magnoliophyta monocots N-fixing claderosids seed plants

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Abietales Acanthales Acerales Acorales Actinidiales Actinostrobales Adoxales Aescul

ales Agavales Aizoales AkanialesAlismatales Alliales Alseuosmiales Alstroemeriales

Altingiales Amaranthales Amaryllidales Amborellales AmbrosialesAmmiales Amomal

es Ancistrocladales Anisophylleales Annonales Anthobolales Apiales Apocynales Apo

nogetonalesAquifoliales Arales Araliales Aralidiales Araucariales Arecales Aristolochi

ales Athrotaxidales Asarales AsclepiadalesAsparagales Asphodelales Asparagales Ast

eliales Atriplicales Aucubales Austrobaileyales Avenales

Balanitales Balanopales Balanophorales Balsaminales Barbeyales Barclayales Batales

Begoniales BerberidalesBerberidopsidales Betulales Biebersteiniales Bignoniales Bix

ales Boraginales Brassicales Brexiales Bromeliales BrunialesBrunoniales Burmannial

es Burserales Butomales Buxales Byblidales

Cactales Callitrichales Calycanthales Calycerales Campanulales Campynematales Can

ellales Cannales CapparalesCaprifoliales Cardiopteridales Carduales Caricales Carle

manniales Caryophyllales Cassiales Casuarinales CelastralesCentrolepidales Cephalot

ales Cephalotaxales Ceratophyllales Cercidiphyllales Chenopodiales Chironiales Chlor

anthalesChrysobalanales Cinchonales Circaeasterales Cistales Citrales Cocosales Col

chicales Columelliales CombretalesCommelinales Connarales Convolvulales Coriarial

es Cornales Corylales Corynocarpales Crassulales CrossosomatalesCucurbitales Cunn

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

40

inghamiales Cunoniales Cupressales Cycadales Cyclanthales Cymodoceales Cynarales

CynomorialesCyperales Cyrillales Cytinales

Daphnales Daphniphyllales Dasypogonales Datiscales Degeneriales Desfontainiales D

iapensiales Didymelales DillenialesDioales Dioncophyllales Dioscoreales Diospyrales

Dipentodontales Dipsacales Droserales

Ebenales Echiales Elaeagnales Elaeocarpales Elatinales Elodeales Emmotales Empetr

ales Ephedrales Ericales EriocaulalesErythropalaless Escalloniales Eucommiales Euph

orbiales Eupomatiales Eupteleales Euryalaless

Fabales Fagales Falcatifoliales Ficales Flacourtiales Flagellariales Fouquieriales Franc

oales Frangulales

Galiales Garryales Geissolomatales Gentianales Geraniales Gesneriales Ginkgoales G

laucidiales Globulariales GnetalesGoodeniales Greyiales Griseliniales Grossulariales

Grubbiales Gunnerales Gyrocarpales Gyrostemonales

Haemodorales Haloragales Hamamelidales Hanguanales Heisteriales Helleborales He

lwingiales Himantandrales HippuridalesHomaliales Hortensiales Huales Huerteales

Hydatellales Hydnorales Hydrangeales Hydrastidales HydrocharitalesHydropeltidales

Hydrostachyales Hypericales Hypoxidales

Icacinales Illiciales Iridales Irvingales Iteales Ixiales

Jasminales Juglandales Julianiales Juncaginales Juncales

Lacistematales Lactoridales Lamiales Lardizabalales Laurales Lecythidales Ledocarpal

es Leitneriales LentibularialesLigustrales Liliales Limnanthales Linales Liriales Loasal

es Lobeliales Loganiales Lonicerales Loranthales LowialesLythrales

Magnoliales Malpighiales Malvales Marathrales Marcgraviales Mayacales Medusagy

nales Medusandrales MelanthialesMelastomatales Meliales Melianthales Meliosmal

es Menispermales Menyanthales Metteniusales MitrastemonalesMiyoshiales Moni

miales Moringales Musales Myricales Myristicales Myrothamnales Myrsinales Myrta

les

Najadales Nandinales Narcissales Nartheciales Nelumbonales Nepenthales Neuradal

es Nitrariales Nolanales NothofagalesNyctaginales Nymphaeales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

41

Ochnales Oenotherales Olacales Oleales Onagrales Oncothecales Opuntiales Orchid

ales Oxalidales

Saxifragales Pandales Pandanales Papaverales Paracryphiales Parasitaxales Paridales

Parnassiales PassifloralesPenaeales Pennantiales Pentaphragmatales Peridiscales P

etiveriales Petrosaviales Phellinales Philydrales PhyllanthalesPhysenales Phytolaccal

es Picramniales Pinales Pinguiculales Piperales Pittosporales Plantaginales Platanale

sPlumbaginales Poales Podocarpales Podophyllales Podostemales Polemoniales Pol

ygalales Polygonales PontederialesPortulacales Posidoniales Potamogetonales Primu

lales Proteales

Quercales Quillajales Quintiniales

Rafflesiales Ranunculales Rapateales Resedales Restionales Rhabdodendrales Rhamn

ales Rhinanthales RhizophoralesRhodorales Rhoipteleales Roridulales Rosales Rouss

eales Rubiales Ruppiales Rutales

Sabiales Salicales Salvadorales Samolales Samydales Sanguisorbales Santalales Sapin

dales Sapotales SarracenialesSaxegotheales Saxifragales Scheuchzeriales Sciadopitya

les Scleranthales Scrophulariales Scyphostegiales SedalesSilenales Simmondsiales S

milacales Solanales Sphenocleales Sphenostemonales Stangeriales Stellariales Stemo

nalesStilbales Stylidiales Styracales Surianales

Taccales Tamales Tamaricales Taxales Taxodiales Tecophilaeales Terebinthales Tern

stroemiales Theales TheligonalesThymelaeales Tiliales Tofieldiales Torricelliales Tov

ariales Tribelales Trilliales Trimeniales Triuridales TrochodendralesTropaeolales Turn

erales Typhales

Ulmales Urticales

Vacciniales Vahliales Vallisneriales Velloziales Veratrales Verbenales Viburnales Vinc

ales Violales Viscales VitalesVochysiales

Welwitschiales Winterales

Xanthorrhoeales Ximeniales Xyridales

Zamiales Zingiberales Zosterales Zygophyllales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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ALPHABETICAL LISTING OF ALL FAMILY NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | K | L | M | N | O | P | Q | R | S | T | U | V | W | X | Y

|Z

Back to Home

In some cases clicking on a family synonym will send you directly to the subfamily in

which the genus of the basionym is to be found

Abietaceae Abolbodaceae Abrophyllaceae Acalyphaceae Acanthaceae [Acanthochla

mydaceae] Acarnaceaee AceraceaeAchariaceae Achatocarpaceae Achradaceae Achy

ranthaceae Acmopylaceae Acoraceae Acristaceae ActaeaceaeActinidiaceae Actinota

ceae Amentotaxaceae Adenogrammaceae Adoxaceae Aegialitidaceae Aegicerataceae

Aegilopaceae Aeginetiaceaee Aegiphilaceae Aesculaceae Aextoxicaceae Agapanthac

eae Agavaceae AgdestidaceaeAgialidaceae Agrimoniaceae Agrostidaceae Ailanthace

ae Aitoniaceae Aizoaceae Akaniaceae Alangiaceae AlchemillaceaeAldrovandaceae A

lismataceae Alliaceae Allioniaceae Allophylaceae Aloaceae Alopecuraceae Alpiniacea

e AlseuosmiaceaeAlsinaceae Alsodeiaceae Alstroemeriaceae Altingiaceae Alzateacea

e Amaranthaceae Amaryllidaceae Amborellaceae Ambrosiaceae Amentotaxaceae A

mmanniaceae Ammiaceae Amomaceae Ampelidopsaceae Amygdalaceae Amyridacea

eAnacardiaceae Anagallidaceae Anarthriaceae Anchusaceae Ancistrocladaceae Andr

omedaceae AndropogonaceaeAndrostachyaceae Androsynaceae Anemarrhenaceae

Anemonaceae Angelicaceae Anisophylleaceae AnnonaceaeAnomochloaceae Anopter

aceae Anrederaceae Anthemidaceae Anthericaceae Anthobolaceae Antidesmataceae

Antirrhinaceae Antoniaceae Aparinaceae Aphanopetalaceae Aphloiaceae Aphyllanth

aceae Apiaceae ApocynaceaeApodanthaceae Aponogetonaceae Aporusaceae Apose

ridaceae Apostasiaceae Aptandraceae AquifoliaceaeAquilariaceae Aquilegiaceae A

raceae Aragoaceae Araliaceae Aralidiaceaee Araucariaceae Arbutaceae Arceuthidace

aeArceuthobiaceae Arctostaphylaceae Arctotidaceae Ardisiaceae Arecaceae Argoph

yllaceae Arisaraceae AristolochiaceaeAristoteliaceae Arjonaceae Armeriaceae Artem

isiaceae Arthrotaxidaceae Artocarpaceae Arundinaceae ArundinellaceaeAsaraceae A

sclepiadaceae Ascyraceaee Asiraceae Asparagaceae Asperulaceae Asphodelaceae As

pidistraceaeeAsteliaceae Asteraceae Asteranthaceae Asteropeiaceae Astragalaceae

Astrocarpaceae AthanasiaceaeAtherospermataceae Atriplicaceae Atropaceae Aucuba

ceae Aurantiaceae Austrobaileyaceae Austrotaxaceae AvenaceaeAverrhoaceae Avet

raceae Avicenniaceae Azaleaceae Azimaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Balanitaceae Balanopaceae Balanophoraceae Balsameaceae Balsaminaceae Bambusa

ceae Barbaceniaceae BarbeuiaceaeBarbeyaceae Barclayaceae Barringtoniaceae Base

llaceae Bataceae Baueraceae Bauhiniaceae Baxteriaceae BegoniaceaeBehniaceae B

elangeraceae Belloniaceae Bembiciaceae Berberidaceae Berberidopsidaceae Berryac

eaee BersamaceaeBertyaceae Berzeliaceae Besleriaceae Betaceae Betulaceae Biebe

rsteiniaceae Bifariaceae Bignoniaceae BischofiaceaeBixaceae Blakeaceae Blandfordia

ceae Blattiaceae Blepharocaryaceae Blitaceae Blyxaceae Boerlagellaceae Bolivaracea

eBombacaceae Bonnetiaceae Bontiaceae Boopidaceae Boraginaceae Borassaceae B

oroniaceae BoryaceaeBotryodendraceae Bougainvilleaceae Boweniaceae Brassicacea

e Brachycaulaceae Bretschneideraceae BrexiaceaeBromeliaceae Brownlowiaceae Br

unelliaceae Bruniaceae Brunoniaceae Brunsvigiaceae Bryoniaceae BuchneraceaeBuci

daceae Buddlejaceae Buglossaceae Bulbocodiaceae Bumeliaceae Bupleuraceae Burc

hardiaceae BurmanniaceaeBurseraceae Butneriaceae Butomaceae Buxaceae Byblida

ceae Byttneriaceae

Cabombaceae Cacaoaceae Cactaceae Caesalpiniaceae Caladiaceae Calamaceae Calce

olariaceae CalectasiaceaeCalendulaceae Callaceae Callicomaceae Calligonaceae Calli

traceae Callitrichaceae Calochortaceae CalophyllaceaeCalthaceae Calycanthaceae Ca

lyceraceae Cambogiaceae Camelliaceae Campanulaceae CampynemataceaeCanacom

yricaceae Canellaceae Cannabaceae Cannaceae Canopodaceae Canotiaceae Cansjera

ceae CapparaceaeCaprariaceae Caprifoliaceae Cardiopteridaceae Carduaceae Carica

ceae Carissaceae Carlemanniaceae CarpinaceaeCarpodetaceae Cartonemataceae Ca

ryocaraceae Caryophyllaceae Casearieaceae Cassiaceae CassipoureaceaeCassythacea

e Castaneaceae Castelaceae Casuarinaceae Catesbaeaceae Cathedraceae Cecropiace

ae CedraceaeCedrelaceae Celastraceae Celosiaceae Celtidaceae Centaureaceae Cen

trolepidaceae Centroplacaceae CepaceaeCephalanthaceae Cephalotaceae Cephalota

xaceae Cerastiaceae Ceratoniaceae Ceratophyllaceae CerberaceaeCercidiphyllaceae

Cercocarpaceae Cercodiaceae Cereaceae Cerinthaceae Ceroxylaceae Cestraceae Cev

alliaceaeChailletiaceae Chamaedoraceae Chamaemoraceae Chamelauciaceae Chaun

ochitonaceae Chelidoniaceae ChelonaceaeChenopodiaceae Chimonanthaceae Chingi

thamnaceae Chionographidaceae Chiranthodendraceae ChironiaceaeChlaenaceae Ch

loanthaceae Chloranthaceae Chloridaceae Chlorogalaceae Chrysobalanaceae Cicerac

eae CichoriaceaeCimicifugaceae Cinchoniaceae Circaeaceae Circaeasteraceae Cissac

eae Cistaceae Citraceae Clematidaceae CleomaceaeClethraceae Clusiaceae Cneorac

eae Cnestidaceae Cnicaceae Cobaeaceae Cochlospermaceae Cocosaceae Coffeaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Colchicaceae Coleogynaceae Columelliaceae Combretaceae Commelinaceae Comocl

adiaceae Compositae CompsoaceaeConantheraceae Connaraceae Conostylidaceae C

onvallariaceae Convolvulaceae Cordiaceae CoreopsidaceaeCoriandraceae Coriariacea

e Coridaceae Corispermaceae Cornaceae Corokiaceae Coronillaceae Correaceae Cor

rigiolaceaeCorsiaceae Corydalaceae Corylaceae Corynocarpaceae Coryphaceae Cost

aceae Cotyledonaceae CoulaceaeCoutariaceae Coutoubiaceae Crassulaceae Crescen

tiaceae Cressaceae Crinaceae Crocaceae CroomiaceaeCrossosomataceae Crotonacea

e Cruciferae Cryptaceae Crypteroniaceae Cryptocorynaceae CryptomeriaceaeCtenol

ophonaceae Cucurbitaceae Cunninghamiaceae Cunoniaceae Cupressaceae Curcumac

eae Curtisiaceae CuscutaceaeCyananthaceae Cyanastraceae Cynanchaceae Cyanella

ceae Cycadaceae Cyclanthaceae CyclantheraceaeCyclocheilaceae Cydoniaceae Cymo

doceaceae Cynaraceae Cynocrambaceae Cynomoriaceae Cyperaceae CyphiaceaeCyp

hocarpaceae Cypripediaceae Cyrillaceae Cyrtandraceae Cyrtanthaceae Cytinaceae

Dacrycarpaceae Dactylanthaceae Damasoniaceae Daphnaceae Daphniphyllaceae Das

ypogonaceae DatiscaceaeDaturaceae Daucaceae Davidiaceae Davidsoniaceae Decais

neaceae Deeringiaceae Degeneriaceae DelphiniaceaeDendrophthoaceae Columelliac

eae Detariaceae Dialypetalanthaceae Dianellaceae Dianthaceae DiapensiaceaeDicha

petalaceae Dichondraceae Diclidantheraceae Dicrastylidiaceae Dictamnaceae Didiere

aceae DidymelaceaeDidymocarpaceae Diegodendraceae Diervillaceae Digitalidaceae

e Dilatridaceae Dilleniaceae Dioaceae DionaeaceaeDioncophyllaceae Dioscoreaceae

Diosmaceae Diospyraceae Dipentodontaceae Diphylleiaceae Diplolaenaceae Dipsaca

ceaeDipterocarpaceae Dirachmaceae Disanthaceae Diselmaceae Dodonaeaceae Do

mbeyaceae Donatiaceae DorsteniaceaeDortmannaceae Doryanthaceae Drabaceae D

racaenaceae Dracontiaceae Drimyidaceae Droseraceae DrosophyllaceaeDryadaceae

Duabangaceae Duckeodendraceae Dulongiaceae Durantaceae Durionaceae Dysphani

aceae

Ebenaceae Eccremidaceae Ecdeiocoleaceae Echiaceae Echinopaceae Ehretiaceae Ela

eagnaceae ElaeocarpaceaeElegiaceae Elatinaceae Ellisiaceae Ellisiophyllaceae Elodea

ceae Elytranthaceae Embeliaceae Emblingiaceae EmmotaceaeEmpetraceae Encepha

lartaceae Engelhardtiaceae Enhalaceae Epacridaceae Ephedraceae Ephemeranthacea

eEpilobiaceae Epimediaceae Eragrostidaceae Eremolepidaceae Eremosynaceae Erica

ceae Erinaceae EriocaulaceaeEriogonaceae Eriospermaceae Erodiaceae Erycibaceae

Eryngiaceae Erysimaceae Erythroniaceae ErythropalaceaeErythrospermaceae Erythro

xylaceae Escalloniaceae Eschscholziaceae Eucomidaceae Eucommiaceae Eucryphiace

aeEuonymaceae Eupatoriaceae Euphorbiaceae Euphrasiaceae Euphroniaceae Eupo

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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matiaceae Eupteleaceae EuryalaceaeEustrephaceae Euthemidaceae Eutocaceae Exb

ucklandiaceae Exocarpaceae

Fabaceae Fagaceae Falcatifoliaceae Ferulaceae Festucaceae Ficaceae Ficoidaceae Fit

zroyaceae FlacourtiaceaeFlagellariaceae Flindersiaceae Foetidiaceae Forestieraceae

Fothergillaceae Fouquieriaceae Fragariaceae FrancoaceaeFrangulaceae Frankeniacea

e Fraxinaceae Fraxinellaceae Freycinetiaceae Fritillariaceae Fuchsiaceae Fumariacea

eFunkiaceae

Gaiadendraceae Galacaceae Galanthaceae Galaxiaceae Galedupaceae Galeniaceae G

aliaceae Garciniaceae GardeniaceaeGardneriaceae Garryaceae Geissolomataceae Ge

itonoplesiaceae Gelsemiaceae Geniostomaceae GentianaceaeGeonomataceae Geosir

idaceae Geraniaceae Gerrardinaceae Gesneriaceae Gethyllidaceae Gilliesiaceae Gina

lloaceaeGinkgoaceae Gisekiaceae Gladiolaceae Glaucidiaceae Glechomaceae Glinace

ae Globulariaceae Gnaphaliaceae GnetaceaeGoetzeaceae Gomortegaceae Gomphiac

eae Gomphrenaceae Gonystylaceae Goodeniaceae Gordoniaceae GouaniaceaeGoup

iaceae Gramineae Gratiolaceae Grewiaceae Greyiaceae Grielaceae Griseliniaceae Gr

onoviaceae GrossulariaceaeGrubbiaceae Guaiacanaceae Guamatelaceae Guettardac

eae Gunneraceae Gustaviaceae Guttiferae GyrocarpaceaeGyrostemonaceae

Hachetteaceae Haemanthaceae Haemodoraceae Halesiaceae Hallieraceae Halocarpa

ceae Halophilaceae HalophytaceaeHaloragaceae Hamamelidaceae Hanguanaceae H

aptanthaceae Harmandaceae Hebenstretiaceae HectorellaceaeHederaceae Hedyosm

aceae Hedyotidaceae Hedysaraceae Heisteriaceae Heleniaceae Heliamphoraceae H

elianthaceaeHelianthemaceae Helichrysaceae Heliconiaceae Helicteraceae Heliotropi

aceae Helleboraceae Heloniadaceae HelosidaceaeHelwingiaceae Hemerocallidaceae

Hemimeridaceae Hemiphylacaceae Henriqueziaceae Henslowiaceae Hermanniaceae

Hernandiaceae Herniariaceae Herreriaceae Hesperocallidaceae Heterantheraceae He

teropyxidaceae HeterostylaceaeHibbertiaceae Hibiscaceae Hilleriaceae Himantandra

ceae Hippocastanaceae Hippocrateaceae HippomanaceaeHippophaeaceae Hippurida

ceae Hirtellaceae Holacanthaceae Homaliaceae Hopkinsiaceae Hoplestigmataceae H

ordeaceaeHornschuchiaceae Hortensiaceae Hortoniaceae Hostaceae Houstoniaceae

Huaceae Huerteaceae HugoniaceaeHumbertiaceae Humiriaceae Hyacinthaceae Hyd

atellaceae Hydnoraceae Hydrangeaceae Hydrastidaceae HydrillaceaeHydrocharitacea

e Hydrocotylaceae Hydrogetonaceae Hydroleaceae Hydropeltidaceae Hydrophylacac

eae HydrophyllaceaeHydrostachyaceae Hymenocardiaceae Hypecoaceae Hyperanth

eraceae Hypericaceae Hypopityaceae HypoxidaceaeHypseocharitaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta

ceae ImperatoriaceaeInocarpaceae Inulaceae Ionidiaceae Iriarteaceae Iridaceae Irvi

ngiaceae Isnardiaceae Isophysidaceae IteaceaeStrasburgeriaceae Ixiaceae Ixioliriace

ae Ixonanthaceae

Jalapaceae Jamboliferaceae Japonoliriaceae Jasionaceae Jasminiaceae Johnsoniaceae

Joinvilleaceae JuglandaceaeJulianiaceae Juncaceae Juncaginaceae Juniperaceae Jus

siaeaceae Justiciaceae

Kadsuraceae Kaliphoraceae Kalmiaceae Kaniaceae Kiggelariaceae Kingdoniaceae Kin

giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae

Krameriaceae

Labiatae Lacandoniaceae Lachenaliaceae Lacistemataceae Lactoridaceae Lactucaceae

Lagerstroemiaceaee LamiaceaeLampsanaceae Lanariaceae Langsdorffiaceae Lantan

aceae Lapageriaceae Lardizabalaceae Lasiopetalaceae LauraceaeLawsoniaceae Laxm

anniaceae Lecythidaceae Ledaceae Ledocarpaceae Leeaceae Leguminosae Leitneriac

eae LemnaceaeLennoaceae Lentibulariaceae Lentiscaceae Leoniaceae Leonticaceae

Lepidobotryaceae LepidocarpaceaeLepidocaryaceae Lepidocerataceae Leptaulaceae

Leptospermaceae Lepturaceae Lepuropetalaceae LeuctenbergiaceaeLibocedraceae

Licaniaceae Ligustraceae Lilaeaceae Liliaceae Limeaceae Limnanthaceae Limnocharit

aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace

ae Linderniaceae Lindleyaceae Linnaeaceae LippayaceaeLiriaceae Liriodendraceae Li

ssocarpaceae Littorellaceae Loasaceae Lobeliaceaee Loganiaceae LomandraceaeLoni

ceraceae Lopeziaceae Lophiocarpaceae Lophiolaceae Lophiraceae Lophophytaceae L

ophopyxidaceae LoranthaceaeLotaceae Lowiaceae Lupulaceaee Luxemburgiaceae Lu

zuriagaceae Lychnidaceae Lyciaceae LyginiaceaeLygodisodeaceae Lysimachiaceae Ly

thraceae

Macarisiaceae Mackinlayaceae Madiaceae Maesaceae Magnoliaceae Malaceae Male

sherbiaceae MalortieaceaeMalpighiaceae Malvaceae Mangiaceae Manicariaceae Ma

paniaceae Marantaceae Marathraceae MarcgraviaceaeMartyniaceae Mastixiaceae

Matricariaceae Maundiaceae Mayacaceae Medeolaceae Medusagynaceae Peridiscac

eaeMelaleucaceae Melampyraceae Melanophyllaceae Melanthiaceae Melastomatac

eae Meliaceae Melianthaceae MelicaceaeMeliosmaceae Melittidaceae Melochiacea

e Memecylaceae Mendonciaceae Menispermaceae Menthaceae MenyanthaceaeMe

nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

47

Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

Nypaceae Nyssaceae

Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

ceae Papilionaceae Pappophoraceae PapyraceaeParacryphiaceae Parasitaxaceae Par

ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

48

hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

49

cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

50

ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 21: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

21

A PHYLOGENETIC TREE OF SEED PLANTS (THE MAIN TREE)

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

22

The Main Tree above is a conservative summary of well supported relationships

between the major clades of extant seed plants Reading - interpreting and

understanding - such a phylogenetic tree is not necessarily easy so reading a tree may

require a bit of practice OHara (eg 1988 1992) provides an accessible introduction to

thinking and talking about history in the context of looking at phylogenetic trees More

recently Krell and Cranston (2004) Crisp and Cook (2005) and others have emphasized

how careful one must be when interpreting and talking about evolution in ladderized

trees in particular and phylogenetic trees in general Gregory (2008) provides a

particularly detailed discussion of this problem In ladderized trees the smaller (in terms

of numbers of terminals) sister taxon is consistently shown on the same side at every

node thus the trees tend to be pectinate like the teeth of a comb however the

horizontal axis that is as a result so evident has no polarity or particular direction The

use of the adjective basal when discussing phylogenetic trees is especially dangerous

(see also Wojciechowski et al 2004 D Soltis et al 2005b) When I use the term and the

context is not otherwise clear I am referring to the pectinations at the base of a

ladderised tree Remember that when talking about sister taxa one can never be basal

to the other or older or younger although members of one can be more derived - in the

sense of having more apomorphies - than the other (but even then one has to be

careful) The words primitive and advanced and lower and higher should

practically never be used especially when talking about taxa Amborellaceae and

Pinaceae are sister to all other flowering plants and Pinales and are basal to the crown

groups of all other flowering plantsPinales respectively but that does not mean they

are thereby primitive Pinaceae in particular have numerous apomorphies The word

plesiomorphic is far less loaded than primitive and can be used to talk about

individual characters

Most trees here have been more or less ladderized as is the one above The

ladderization may be imperfect for example you can see that on the Main Tree

asterids with ten orders follow rosids and relatives which have seventeen although

this is in part because of the currently unresolved nature of relationships between the

main clades of core eudicots (this is likely to change soon) The Main Tree could have

been drawn with Amborella Acorus or a host of other taxa at the far right without

offending any relationships phylogenetic trees are like mobiles the only fixed points

being the nodes Despite the problems just mentioned pectination interpreted

carefully has its value and it is not an arbitrary process As one reads the terminals of a

pectinate tree from left to right adjacent terminals will be separated by apomorphies

that will cumulate in an appropriate fashion When thinking of a book or a herbarium

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

23

sequence (see Haston et al 2007) this is of some value Nymphaeales and

Austrobaileyales are here adjacent on the tree but they could be separated by

hundreds of families in the sequence merely by rotating only the node from which

Nymphaeales and their sister group (all other angiosperms minus about 75 species)

arise If adjacent in a book or herbarium or as in these summary pages then it is

relatively easy to relate their characters both apomorphic and plesiomorphic but if

separated by hundreds of pages or two floors in a large building then it is less easy to

get anything from the sequence (Since all orders in the pages of the main part of this

site are preceded by the apomorphies of all nodes immediately below them in the seed

plant phylogeny and because of the linkages that have been built in to the site this

problem is much less serious there there is no sequence) So the particular sequence of

all seed plants that is based on a phylogenetic classification but is to be used in herbaria

or any other place where taxa have to be arranged linearly as in this part of APweb

will be one that maximizes the number of taxa that are both successive branches of the

tree and placed successively in the sequence Since specimens are generally filed under

families the outline of a new family sequence for arranging herbaria and books can now

be suggested (Haston et al 2007) and such a standardized sequence will further help

teaching and learning about plants

ON THINKING ABOUT APOMORPHIES

Identifying apomorphies is important because understanding the evolution of

morphology in the broad sense - ie including anatomy chemistry etc - in the context

of diversification is one of our major goals For identifying apomorphies several

preconditions must be met One needs to have an accurate well supported phylogeny

one has to have examined the right taxa both from the point of view of morphology and

molecules one has to have coded the characters right (ie delimited states

appropriately) and one has to use an appropriate model of evolution when fitting the

variation to the tree (see eg Omland 1999 Stevens 2006b) If the distributions of

apomorphies below are compared with those in other studies (eg Turgeon et al 2001

Bremer et al 2001 Endress 2001 Albach et al 2001a Judd amp Olmstead 2004 D Soltis

et al 2005b Zhang et al 2006 Judd et al 2007) the reader may find a number of

differences Although I have integrated such studies as far as is possible there are five

reasons that link to the preconditions just mentioned why there may be differences

where features are placed on trees

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

24

1 I may not have found all the information about a particular character there may be

disagreement over its interpretation or I have added information

2 The sampling of nearly all molecular studies is very incomplete (see Salisbury amp Kim

2001 for problems caused by sampling) indeed it is commonplace to decry the

incompleteness of molecular sampling But not only is the sampling in molecular studies

often less than we might wish that of the morphological and chemical characters whose

evolution we are interested in understanding is also often very poor So for many

anatomical chemical and embryological characters that are confidently said to

characterise families and other groups we all too often have no idea if those characters

are applicable to the whole clade or just to a subgroup within that clade However as

our sampling improves we can locate changes on the tree more precisely if in

unexpected positions Thus is is sometimes suggested that Ericaceae have ellagic acid

(Soltis et al 2005b) in fact the little evidence we have indicates that only a small clade

of some 80 species has ellagic acid and the rest of the family - itself alone about one

third of all Ericales - does not Similarly Ericaceae are noted for their

ectendomycorrhizal associations but Enkianthus sister to all other Ericaceae

apparently lacks these (Abe 2005) Thus these mycorrhizae and their associated

distinctive hair roots are unlikely to be apomorphies of Ericaceae - however it appears

that other families near Ericaceae may also have ectendomycorrhizae (see Asai 1934)

3 I am fitting characters to a very conservative tree with many polytomies although the

nodes that are utilised are for the most part strongly supported polytomies make the

optimisation of characters that is the assigment of character state change to a

particular node on the tree notably difficult (eg Madison amp Madison 2002) In nearly

all studies of the evolution of characters distributions of characters are optimised on a

more or less fully resolved tree and the construction of supertrees may yield yet more

detailed hypotheses of relationships (for literature on supertrees see Cotton amp

Wilkinson 2007 2008) Of course some nodes on such fully resolved trees andor

supertrees may have little support and optimisations of characters on such trees may

carry correspondingly little conviction Even parts of some of the trees used here have

poor support eg relationships within aquatic Alismatales etc although I indicate

when this is the case

4 Exactly how one goes about optimising a character on a tree is critically important

Even using simple parsimony optimisations (ACCTRAN or DELTRAN ACCelerated

TRANsitions or DELayed TRANsitions) the position of synapomorphies on trees - and

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

25

hence our ideas of evolution - may differ and this is still more true if one uses maximum

likelihood or Bayesian methods and makes apparently reasonable suggestions about

weighting gains over losses (eg Donoghue amp Ackerley 1996 Cunningham et al 1998

Omland 1997 1999 Ree amp Donoghue 1999 Polly 2001 Webster amp Purvis 2001

Ronquist 2004 Crisp amp Cook 2005) Sannier et al (2007) show how in in Arecaceae that

where on a phylogenetic tree one might peg changes in microsporogenesis will depend

on the methods one uses to do this and Pedersen et al (2007) discuss the sometimes

very substantial effect of node support on the posterior probabilities of ancestral

character states Here I use parsimony optimization not always as explicit as it might be

but I have often indicated where there are particularly important uncertainties as to the

positions of particular character changes on the tree

5 Finally although I have paid quite a lot of attention to the delimitation of the

character states that make up all the characterizations I have not spent enough time on

this critical operation If we are interested in understanding evolution then fitting the

basic variation - not character states - to a tree in principle allows greater flexibility in

understanding morphology in the context of local phylogenies (see also Stevens 2000

Endress 2005c) However many character states used here are delimited globally that

is they are circumscribed in the context of the variation shown by individual characters

across all angiosperms andor in the context of classic ideas of character evolution

Character states often have arbitrary limits and serve best to communicate

information whether they are in fact suitable for either phylogenetic analysis or

understanding evolution are separate issues Studies have rather unsurprisingly

perhaps but importantly shown that dividing the one character into different sets of

states may yield differing ideas of evolution of that character (eg Lamb Frye amp Kron

2003 Hibbett 2004) When looking at trees on which character states are optimised

one should bear in mind the problems surrounding the delimitation of states (eg

Stevens 2000 2006b) and the danger of using pollen or other types - constructs

based on many characters that vary independently but which effectively get lost in

these types thus Blackmore et al (2009) decomposed the pollen types in Asteraceae

into 52 characters

I use Remanes three main criteria of homology or better similarity when

determining the basic similarity of structures on different organisms (see Remane 1952)

These criteria are special properties position and intermediates Special properties

include anatomical or chemical characters gene expression data etc Position refers

to the position of an organ with respect to landmarks on the plant Although plants are

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

26

plants and landmarks are not as fixed as one might like trying to work out the relative

positions of parts is a good way of understanding morphology so long as one does not

try to out-do Procrustes Intermediates include intermediates found on the same

plant or on different plants Intermediates may observed during development or be

adult structures Thus one may find changes during development which suggest what

the nature of a particular structure is and very different stuctures on different plants

may turn out to be similar early in development Or one can simply compare different

taxa and see that structures that appear to be very different when comparing two

different taxa can be linked morphologically by looking at other taxa

Nevertheless the use of these criteria may not yield an unambiguous answer as to

what a structure is even given a solid phylogeny and an improved understanding of

development (see Jaramillo amp Kramer 2007 for a useful discussion) As Endress (2005c)

observed a number of features - position function development shape anatomy

histology gene activity and relationships to other taxa that clearly have petals - can be

used to distinguish a petal (for example) from other floral structures if a petal does not

have one of these features is it thereby not a petal Thus Maturen et al (2005) recently

found that floral organ diversity genes (B and C) were expressed in the large white

inflorescence bracts of Cornus (see also Costa et al 2005) Peney et al (2005) noted that

not all monosulcate pollen grains in monocots have the same developmental pathway

and that as a result such pollen might not have the same ancestral state Reeves and

Olmstead (2003) suggested that the genetic mechanisms causing monosymmetry in

Lamiales and Solanales were different and Serna and Martin (2006) described similar

problems with the development of hairs in Arabidopsis when compared with that of

hairs in Antirrhinum and Solanaceae Indeed as one perhaps might expect delimitation

of states and characters does not necessarily become easier with increasing knowledge

of development etc Thus Buzgo et al (2004) Matthews and Endress (2005) and others

have shown how hard it can be to distinguish between eg prophylls and other floral

structures as their behaviour is studied during the course of the development of the

flower Similarly at what concentration is a particular secondary metabolite deemed to

be present (Waterman (2007)

The validity of the approach used here that of fitting morphological variation to a

largely molecular-based tree may be questioned However I think it rather unlikely that

well-supported molecular branches will be overturned by morphological data Indeed

analyses of morphological data alone do provide support for many of the clades evident

in molecule-only analyses and analyses with morphological and molecular data

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

27

together may lead to increased support for clades (eg Hufford 1992 Nandi et al 1998

[but adding morphological data reduces support for a number of critical clades too]

Doyle amp Endress 2000 however in none of these papers is the use of morphology

without ambiguity) It is unfortunately clear that the use of morphology alone may not

suggest problems in the phylogenetic placement of taxa that later turn out to have been

wrongly included (eg Zhang et al 1992) Thus I have been wary of putting much weight

on clades that have only morphological support but note that I have treated molecule-

based clades with low bootstrap or jacknife support values (esp below 70) or low

posterior probabilies (below 095) likewise Although I may have been mistaken in

placing so much emphasis on molecular data in terms of providing the basic

phylogenetic framework for angiosperms morphological and molecular data are only

very rarely in irreconcilably strong conflict There are indeed a few places where the

conflict seems extreme These include the relative positions of the Monimiaceae and

Hernandiaceae (Laurales) the position of Hanguanaceae (Commelinales [as here] or

Zingiberales) and of Triplostegia (is it in Dipsacaceae or Valerianaceae - see

Dipsacales) Fossils are unlikely to affect the topologies of the trees presented here but

see below for their importance in understanding morphological evolution in general and

the evolution of angiosperms in particular Nevertheless some largely reject the idea

that trees based on molecular data alone can recover phylogenetic relationships

especially when branching points are old and prefer to used trees based on analysis of

morphological data including those taken from fossils (Hilton amp Bateman 2006 Farjon

2007)

All in all however the extent of the congruence between morphological and

molecular data is impressive and heartening and many clades can be characterised

morphologically It seemed in 1998 that there were no unambiguous morphological

synapomorphies for angiosperm orders (K Bremer 2000) and this is still true if by

unambiguous is meant non-homoplasious However many orders and other clades

have synapomorphies even if these may be indistinguishable at least at the current

level of morphological and developmental knowledge from parallel occurences

elsewhere As our morphological knowledge increases so too does the number of

apomorphies

Endress and Matthews (2006a) emphasize the importance of tendencies and

developmental constraints when thinking about characters of clades within the rosids -

many characters or character combinations occur in a rather sporadic fashion within

the clade and apparently notably less frequently outside it There are a number of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

28

examples of tendencies here and in the individual order pages such as the features

enclosed by parentheses in groups above the level of family eg see (cuticular waxes

as aggregated rodlets) for the commelinids and the discussion of the distribution of

polyandry that is flowers with numerous stamens in the asterid I + II groups etc

Tendencies may involve more than single characters When thinking of the

associations of features like integument thickness and vascularization of the integument

(Endress amp Matthews 2006a) one can imagine fairly simple developmental

preconditions being operative As Endress and Matthews (2006a) note it is difficult to

imagine vascular tissue developing in an integument only two cells thick Changes here

would be loosely correlated if morphologically linked However the strongly correlated

changes noted by Givnish et al (2005) are ecologically linked but are presumably

morphologicallydevelopmentally independent When there seem to be characters

evolving more or less together tests can be carried out to see if the changes are

concentrated on certain branches of the tree (eg Maddison 1990 Sanderson 1991

Maddison amp Maddison 2000) Such tests have rarely been carried out at higher levels in

angiosperms

One of the most striking examples of a tendency is the distribution of N-fixation

restricted as it is to a monophyletic group of four clades although it has arisen seven (or

perhaps several more) times independently within the N-fixing clade and several

members of two quite different kinds of bacteria are involved (eg Clawson et al 2004

Elliott et al 2007 Sprent amp James 2007) Other examples of work that bears on the issue

of tendencies include the findings that flowers of polysymmetrical Arabidopsis have

genes like TCP1 that are expressed asymmetrically during early development and TCP1

is a probable orthologue of the well-known CYC gene of Antirrhinum that is involved in

the development of monosymmetric slowers there (Cubas et al 2001 Costa et al 2005

etc) Parallelism might build on this underlying morphologically cryptic monosymmetry

even if details of the genetic mechanisms causing the monosymmetry evident in

particular groups may be different (Reeves amp Olmstead 2003 Cubas 2004 see above)

Similarly there have been several recent suggestions that the capability to synthesise a

particular metabolite may be switched off but not lost and so can sometimes be

reacquired (eg Wink amp Witte 1983 Wink 2003 Liscombe et al 2005 Larsson 2007

Waterman 2007) Hence perhaps the rather spotty distribution of many secondary

metabolites like ellagic acid the indole alkaloid camptothecin iridoids etc when

considered in the context of phylogenies As a non-botanical example - but a rather nice

one - Salwini-Plawen and Mayr (1961) suggested some time ago that there has been

considerable parallelism (40-65 or more independent origins) in the evolution of eyes in

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

29

metazoans However Pax 6 seems to be a master control gene that is involved in eye

formation perhaps throughout all bilateralians (eg Gehring amp Ikeo 1999 Erwin amp

Davidson 2002) which changes the nature of the problem Note however that in

general our invocation of evolutionary tendencies to explain the patterns of

distributions of characters is really an admission that we do not understand those

patterns

As we find out more about variation we will probably find fewer and fewer features

constant throughout a group Most unqualified statements of presence and absence

should properly be qualified as usually present or usually absent if one is thinking of

the characterisations on the order pages as encompassing the total variation within a

clade Thus Pistia alone among monocots as so far known has sieve tube plastids with

starch grains not protein crystals However this does not affect the fact that sieve tube

plastids with cuneate protein crystals and lacking starch grains are an apomorphy for

monocots Furthermore fossils in a number of cases suggest character combinations

unknown in extant taxa as may be seen in the discussions of Fagaceae Platanaceae

Iteaceae Calycanthaceae etc Confusing the issue there may be questions as to where

exactly on the tree a particular fossil is to be placed (eg see Nymphaeaceae

Calycanthaceae Archaefructus etc)

To summarize given our current understandings of both phylogenies and

characters evolution of some characters in which we are interested seems very labile

(see eg D Soltis et al 2005b Endress and Matthews 2006a Stevens 2006b Ekman et

al 2008 for a good recent study) and I have been cautious when talking about character

evolution Much effort must continue to be spent in summarizing characters of well-

established clades at all levels providing features by which they may be recognized and

signaling synapomorphies Remember that (1) the basic morphological anatomical and

chemical knowledge of many critical taxa is woefully incomplete (2) different

assumptions about character evolution may greatly affect the position of

synapomorphies on trees (3) in many cases relationships within and between many

groups are too uncertain at present to worry very much about synapomorphies and

(4) we must be clear about what we do and do not not know As mentioned above I

have indicated in a number of places where there are particularly important

uncertainties as to where characters should be placed on the tree Nevertheless it is a

relatively easy matter to update notes such as these and it can be a simple matter to

incorporate new data on characters that have never before been considered in the

context of a tree Much basic - and unfortunately perhaps unfashionable - work must be

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

30

carried to clarify the distribution of morphological anatomical and chemical characters

for examples of what can to be done see the work of P K Endress and collaborators

(floral morphology and development) and S R Jensen and collaborators (iridoids) But

acquiring information about nodal anatomy cell and tissue distribution nodal anatomy

and the like is not difficult at all and should be generally encouraged

But all this is merely a necesary prelude to the understanding of evolution For this

one needs to know a lot more including dating the phylogeny understanding the

function(s) of characters factoring in relevant aspects of the palaeoenvironment in

which an apomorphy first appeared etc

SUMMARY OF THE SYSTEM

Below is a formalised summary of the relationships within orders of the families of seed

plants There are a few families that are not recognised even as options in APG II and

vice versa as well as a few extra orders The families to be recognised in the most

recent edition of Mabberleys The Plant Book (Mabberley 2008) are also largely

consistent with those below But all differences are trivial and will - I hope - eventually

disappear

Square brackets - [] - enclose clades the plus sign - + - designates sister taxa a

comma - - denotes part of a polytomy and quotation marks - - denotes a

paraphyletic group

SEED PLANTS

GYMNOSPERMS

Cycadales

Cycadaceae + Zamiaceae

Ginkgoales

Ginkgoaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

31

Pinales

Pinaceae [[Araucariaceae [Podocarpaceae + Phyllocladaceae]] [Sciadopityaceae

[Taxaceae + Cupressaceae]]]

Gnetales

Ephedraceae [Gnetaceae + Welwitschiaceae]

ANGIOSPERMSFLOWERING PLANTS

Amborellales

Amborellaceae

Nymphaeales

Hydatellaceae [Cabombaceae + Nymphaeaceae]

Austrobaileyales

Austrobaileyaceae [Schisandraceae + Trimeniaceae]

Chloranthales

Chloranthaceae

MAGNOLIIDS

Magnoliales

Myristicaceae [Magnoliaceae [[Himantandraceae + Degeneriaceae] [Eupomatiaceae +

Annonaceae]]]

Laurales

Calycanthaceae [[Siparunaceae [Gomortegaceae + Atherospermataceae]] [Monimiaceae

[Hernandiaceae + Lauraceae]]]

Canellales

Canellaceae + Winteraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

32

Piperales

[Hydnoraceae + Aristolochiaceae] [Piperaceae + Saururaceae]

MONOCOTS

Acorales

Acoraceae

Alismatales

Araceae [Tofieldiaceae [[Alismataceae [Hydrocharitaceae + Butomaceae]]

[Scheuchzeriaceae [Aponogetonaceae [Juncaginaceae [Maundiaceae [[Posidoniaceae

[Ruppiaceae + Cymodoceaceae]] [Zosteraceae + Potamogetonaceae]]]]]]]

Petrosaviales

Petrosaviaceae

Dioscoreales

Nartheciaceae [[Taccaceae + Thismiaceae] [Burmanniaceae + Dioscoreaceae]]

Pandanales

Velloziaceae Triuridaceae Stemonaceae [Pandanaceae + Cyclanthaceae]

Liliales

Corsiaceae [Campynemataceae [Petermanniaceae [Colchicaceae + Alstroemeriaceae]]

Melanthiaceae [[Philesiaceae + Rhipogonaceae] [Smilacaceae + Liliaceae]]]

Asparagales

Orchidaceae [[Boryaceae [Blandfordiaceae [Lanariaceae [Asteliaceae + Hypoxidaceae]]]]

[[Ixioliriaceae + Tecophilaeaceae] [Doryanthaceae [Iridaceae [Xeronemataceae

[Xanthorrhoeaceae [Alliaceae + Asparagaceae]]]]]]]

COMMELINIDS

Unplaced

Dasypogonaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

33

Arecales

Arecaceae

Poales

[Typhaceae + Bromeliaceae] [Rapateaceae [[Thurniaceae [Juncaceae + Cyperaceae]]

[[Mayacaceae [Eriocaulaceae + Xyridaceae]] [Flagellariaceae [Anarthriaceae

[Centrolepidaceae + Restionaceae]]] [Joinvilleaceae [Ecdeiocoleaceae + Poaceae]]]]]]]

Commelinales

[Commelinaceae + Hanguanaceae] [Philydraceae [Haemodoraceae + Pontederiaceae]]

Zingiberales

Musaceae [Strelitziaceae + Lowiaceae] Heliconiaceae [[Cannaceae + Marantaceae]

[Costaceae + Zingiberaceae]]

Ceratophyllales

Ceratophyllaceae

EUDICOTS

Ranunculales

Eupteleaceae [Papaveraceae [[[Lardizabalaceae + Circaeasteraceae] [Menispermaceae

[Berberidaceae + Ranunculaceae]]]]

Sabiales

Sabiaceae

Proteales

Nelumbonaceae [Platanaceae + Proteaceae]

Trochodendrales

Trochodendraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

34

Buxales

Haptanthaceae [Buxaceae + Didymelaceae]

CORE EUDICOTS

Gunnerales

Gunneraceae + Myrothamnaceae

Dilleniales

Dilleniaceae

Saxifragales

Peridiscaceae [[Paeoniaceae [Altingiaceae [Hamamelidaceae [Cercidiphyllaceae +

Daphniphyllaceae]]]] [[Crassulaceae [Aphanopetalaceae [Tetracarpaeaceae

[Penthoraceae + Haloragaceae]]]] [Iteaceae [Grossulariaceae + Saxifragaceae]]]]

Cynomoriaceae unplaced

Vitales

Vitaceae

ROSIDS

FABIDROSID I

Zygophyllales

Krameriaceae + Zygophyllaceae

Celastrales

Lepidobotryaceae + Celastraceae

Oxalidales

Huaceae [[Connaraceae + Oxalidaceae] [Cunoniaceae [Elaeocarpaceae [Brunelliaceae +

Cephalotaceae]]]]

Malpighiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

35

[Achariaceae [Goupiaceae [Violaceae + Passifloraceae] [Lacistemataceae Salicaceae]]

[Lophopyxidaceae Putranjivaceae] [Ctenolophonaceae [Erythroxylaceae +

Rhizophoraceae]] Linaceae Ixonanthaceae Humiriaceae Irvingiaceae

Centroplacaceae Pandaceae [[Bonnetiaceae + Clusiaceae] [Calophyllaceae

[Hypericaceae + Podostemaceae]]] [Malpighiaceae + Elatinaceae] Ochnaceae

[Peraceae [Rafflesiaceae + Euphorbiaceae]] [Phyllanthaceae + Picrodendraceae]

[Balanopaceae [[Trigoniaceae + Dichapetalaceae] [Chrysobalanaceae +

Euphroniaceae]]] Caryocaraceae

N-FIXING CLADE

Fabales

Quillajaceae [Fabaceae [Polygalaceae + Surianaceae]]

Rosales

Rosaceae [[Barbeyaceae [Elaeagnaceae Dirachmaceae Rhamnaceae]] [Ulmaceae

[Cannabaceae [Moraceae + Urticaceae]]]]

Cucurbitales

Anisophylleaceae [[Corynocarpaceae + Coriariaceae] [Cucurbitaceae [Tetramelaceae

[Datiscaceae + Begoniaceae]]]] Apodanthaceae

Fagales

Nothofagaceae [Fagaceae [[Myricaceae + Juglandaceae] [Casuarinaceae

[Ticodendraceae + Betulaceae]]]]

MALVIDROSID II

pgtGeraniales

Geraniaceae [[Melianthaceae + Francoaceae] [Vivianaceae + Ledocarpaceae]]

Myrtales

Combretaceae [[Onagraceae + Lythraceae] [[Vochysiaceae + Myrtaceae]

[Melastomataceae [Crypteroniaceae [Alzateaceae + Penaeaceae]]]]]

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

36

Crossosomatales

[Staphyleaceae [Guamatelaceae [Crossosomataceae + Stachyuraceae]]] [Aphloiaceae

[Geissolomataceae + Strasburgeriaceae]]

Picramniales

Picramniaceae

Sapindales

Biebersteiniaceae Nitrariaceae [Kirkiaceae [Anacardiaceae + Burseraceae]]

[Sapindaceae [Simaroubaceae Rutaceae Meliaceae]]

Huerteales

Gerradinaceae [Dipentodontaceae + Tapisciaceae]

Malvales

Neuradaceae [Thymelaeaceae [Sphaerosepalaceae Bixaceae [Cistaceae

[Sarcolaenaceae + Dipterocarpaceae]] [Cytinaceae + Muntingiaceae] Malvaceae]]

Brassicales

[Akaniaceae + Tropaeolaceae] [[Moringaceae + Caricaceae] [Setchellanthaceae

[Limnanthaceae [[Koeberliniaceae [Bataceae + Salvadoraceae]] [Emblingiaceae

[Pentadiplandraceae [Gyrostemonaceae Resedaceae Stixis and relatives] Tovariaceae

[Capparaceae [Cleomaceae + Brassicaceae]]]]]]]]

Berberidopsidales

Aextoxicaceae + Berberidopsidaceae

Santalales

Erythropalaceae Olacaceae [[Loranthaceae [Misodendraceae + Schoepfiaceae]]

[Opiliaceae + Santalaceae]] Balanophoraceae unplaced

Caryophyllales

[[Droseraceae [Nepenthaceae [Drosophyllaceae [Ancistrocladaceae +

Dioncophylleaceae]]]]] [[Frankeniaceae + Tamaricaceae] [Polygonaceae +

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

37

Plumbaginaceae]]] [Rhabdodendraceae [Simmondsiaceae [[Asteropeiaceae +

Physenaceae] [[Caryophyllaceae [Achatocarpaceae + Amaranthaceae]]

[Stegnospermataceae [Limeaceae [[Lophocarpaceae [Barbeuiaceae [Aizoaceae

[Sarcobataceae Phytolaccaceae Nyctaginaceae]]]] [Molluginaceae [Halophytaceae

Montiaceae Basellaceae Didiereaceae [Talinaceae [Portulacaceae [Anacampseros etc +

Cactaceae]]]]]]]]]]]]]

ASTERIDS

Cornales

[Cornaceae + Nyssaceae] [Hydrangeaceae + Loasaceae] Hydrostachyaceae [Curtisiaceae

+ Grubbiaceae]

Ericales

[Balsaminaceae [Marcgraviaceae + Tetrameristaceae]] [[Polemoniaceae +

Fouquieraceae] Lecythidaceae [[Sladeniaceae + Pentaphylacaceae] [Sapotaceae

[Ebenaceae [Maesaceae [Theophrastaceae [Myrsinaceae + Primulaceae]]]]]

[Mitrastemonaceae Theaceae [Symplocaceae [Styracaceae + Diapensiaceae]]

[[Sarraceniaceae [Roridulaceae + Actinidiaceae] [Clethraceae [Cyrillaceae +

Ericaceae]]]]]]

LAMIIDASTERID I

Unplaced

Metteniusaceae Oncothecaceae Icacinaceae and some unplaced ex-Icacinaceae

Garryales

Garryaceae + Eucommiaceae

Unplaced

Boraginaceae Vahliaceae

Gentianales

Rubiaceae [Gentianaceae [Loganiaceae [Gelsemiaceae + Apocynaceae]]]

Lamiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

38

Plocospermataceae [Oleaceae [Tetrachondraceae [Calceolariaceae

[Gesneriaceae Peltanthera [Pedaliaceae Carlemanniaceae Martyniaceae Stilbaceae

Plantaginaceae Linderniaceae Bignoniaceae Verbenaceae Lamiaceae Paulowniaceae

Schlegeliaceae Thomandersiaceae Phrymaceae Rehmannia Acanthaceae

Scrophulariaceae Orobanchaceae Byblidaceae Lentibulariaceae]]]]]

Solanales

[Montiniaceae [Sphenocleaceae + Hydroleaceae]] [Convolvulaceae + Solanaceae]

CAMPANULIDASTERID II

Aquifoliales

[[Cardiopteridaceae + Stemonuraceae] [Aquifoliaceae [Helwingiaceae +

Phyllonomaceae]]

Asterales

[Rousseaceae + Campanulaceae] Pentaphragmataceae [[Alseuosmiaceae [Phellinaceae

+ Argophyllaceae]] Stylidiaceae [Menyanthaceae [Goodeniaceae [Calyceraceae +

Asteraceae]]]]

Escalloniales

Escalloniaceae

Bruniales

[Bruniaceae + Columelliaceae]

Apiales

Pennantiaceae [Torricelliaceae [Griseliniaceae [Pittosporaceae [Araliaceae

[Myodocarpaceae + Apiaceae]]]]]

Paracryphiales

Paracryphiaceae

Dipsacales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

39

Adoxaceae [Diervillaceae [Caprifoliaceae [Linnaeaceae [Morinaceae [Dipsacaceae +

Valerianaceae]]]]]

LINKS TO ORDERS AND FAMILIES

[Back to Top]

MAIN GROUPINGS

Asterids commelinids core eudicots asterid 1 asterid

2 eudicots gymnosperms Magnoliophyta monocots N-fixing claderosids seed plants

ALPHABETICAL LISTING OF ALL ORDINAL NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | L | M | N | O | P | Q | R | S | T | U | V | W | X | Z

Abietales Acanthales Acerales Acorales Actinidiales Actinostrobales Adoxales Aescul

ales Agavales Aizoales AkanialesAlismatales Alliales Alseuosmiales Alstroemeriales

Altingiales Amaranthales Amaryllidales Amborellales AmbrosialesAmmiales Amomal

es Ancistrocladales Anisophylleales Annonales Anthobolales Apiales Apocynales Apo

nogetonalesAquifoliales Arales Araliales Aralidiales Araucariales Arecales Aristolochi

ales Athrotaxidales Asarales AsclepiadalesAsparagales Asphodelales Asparagales Ast

eliales Atriplicales Aucubales Austrobaileyales Avenales

Balanitales Balanopales Balanophorales Balsaminales Barbeyales Barclayales Batales

Begoniales BerberidalesBerberidopsidales Betulales Biebersteiniales Bignoniales Bix

ales Boraginales Brassicales Brexiales Bromeliales BrunialesBrunoniales Burmannial

es Burserales Butomales Buxales Byblidales

Cactales Callitrichales Calycanthales Calycerales Campanulales Campynematales Can

ellales Cannales CapparalesCaprifoliales Cardiopteridales Carduales Caricales Carle

manniales Caryophyllales Cassiales Casuarinales CelastralesCentrolepidales Cephalot

ales Cephalotaxales Ceratophyllales Cercidiphyllales Chenopodiales Chironiales Chlor

anthalesChrysobalanales Cinchonales Circaeasterales Cistales Citrales Cocosales Col

chicales Columelliales CombretalesCommelinales Connarales Convolvulales Coriarial

es Cornales Corylales Corynocarpales Crassulales CrossosomatalesCucurbitales Cunn

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

40

inghamiales Cunoniales Cupressales Cycadales Cyclanthales Cymodoceales Cynarales

CynomorialesCyperales Cyrillales Cytinales

Daphnales Daphniphyllales Dasypogonales Datiscales Degeneriales Desfontainiales D

iapensiales Didymelales DillenialesDioales Dioncophyllales Dioscoreales Diospyrales

Dipentodontales Dipsacales Droserales

Ebenales Echiales Elaeagnales Elaeocarpales Elatinales Elodeales Emmotales Empetr

ales Ephedrales Ericales EriocaulalesErythropalaless Escalloniales Eucommiales Euph

orbiales Eupomatiales Eupteleales Euryalaless

Fabales Fagales Falcatifoliales Ficales Flacourtiales Flagellariales Fouquieriales Franc

oales Frangulales

Galiales Garryales Geissolomatales Gentianales Geraniales Gesneriales Ginkgoales G

laucidiales Globulariales GnetalesGoodeniales Greyiales Griseliniales Grossulariales

Grubbiales Gunnerales Gyrocarpales Gyrostemonales

Haemodorales Haloragales Hamamelidales Hanguanales Heisteriales Helleborales He

lwingiales Himantandrales HippuridalesHomaliales Hortensiales Huales Huerteales

Hydatellales Hydnorales Hydrangeales Hydrastidales HydrocharitalesHydropeltidales

Hydrostachyales Hypericales Hypoxidales

Icacinales Illiciales Iridales Irvingales Iteales Ixiales

Jasminales Juglandales Julianiales Juncaginales Juncales

Lacistematales Lactoridales Lamiales Lardizabalales Laurales Lecythidales Ledocarpal

es Leitneriales LentibularialesLigustrales Liliales Limnanthales Linales Liriales Loasal

es Lobeliales Loganiales Lonicerales Loranthales LowialesLythrales

Magnoliales Malpighiales Malvales Marathrales Marcgraviales Mayacales Medusagy

nales Medusandrales MelanthialesMelastomatales Meliales Melianthales Meliosmal

es Menispermales Menyanthales Metteniusales MitrastemonalesMiyoshiales Moni

miales Moringales Musales Myricales Myristicales Myrothamnales Myrsinales Myrta

les

Najadales Nandinales Narcissales Nartheciales Nelumbonales Nepenthales Neuradal

es Nitrariales Nolanales NothofagalesNyctaginales Nymphaeales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Ochnales Oenotherales Olacales Oleales Onagrales Oncothecales Opuntiales Orchid

ales Oxalidales

Saxifragales Pandales Pandanales Papaverales Paracryphiales Parasitaxales Paridales

Parnassiales PassifloralesPenaeales Pennantiales Pentaphragmatales Peridiscales P

etiveriales Petrosaviales Phellinales Philydrales PhyllanthalesPhysenales Phytolaccal

es Picramniales Pinales Pinguiculales Piperales Pittosporales Plantaginales Platanale

sPlumbaginales Poales Podocarpales Podophyllales Podostemales Polemoniales Pol

ygalales Polygonales PontederialesPortulacales Posidoniales Potamogetonales Primu

lales Proteales

Quercales Quillajales Quintiniales

Rafflesiales Ranunculales Rapateales Resedales Restionales Rhabdodendrales Rhamn

ales Rhinanthales RhizophoralesRhodorales Rhoipteleales Roridulales Rosales Rouss

eales Rubiales Ruppiales Rutales

Sabiales Salicales Salvadorales Samolales Samydales Sanguisorbales Santalales Sapin

dales Sapotales SarracenialesSaxegotheales Saxifragales Scheuchzeriales Sciadopitya

les Scleranthales Scrophulariales Scyphostegiales SedalesSilenales Simmondsiales S

milacales Solanales Sphenocleales Sphenostemonales Stangeriales Stellariales Stemo

nalesStilbales Stylidiales Styracales Surianales

Taccales Tamales Tamaricales Taxales Taxodiales Tecophilaeales Terebinthales Tern

stroemiales Theales TheligonalesThymelaeales Tiliales Tofieldiales Torricelliales Tov

ariales Tribelales Trilliales Trimeniales Triuridales TrochodendralesTropaeolales Turn

erales Typhales

Ulmales Urticales

Vacciniales Vahliales Vallisneriales Velloziales Veratrales Verbenales Viburnales Vinc

ales Violales Viscales VitalesVochysiales

Welwitschiales Winterales

Xanthorrhoeales Ximeniales Xyridales

Zamiales Zingiberales Zosterales Zygophyllales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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ALPHABETICAL LISTING OF ALL FAMILY NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | K | L | M | N | O | P | Q | R | S | T | U | V | W | X | Y

|Z

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In some cases clicking on a family synonym will send you directly to the subfamily in

which the genus of the basionym is to be found

Abietaceae Abolbodaceae Abrophyllaceae Acalyphaceae Acanthaceae [Acanthochla

mydaceae] Acarnaceaee AceraceaeAchariaceae Achatocarpaceae Achradaceae Achy

ranthaceae Acmopylaceae Acoraceae Acristaceae ActaeaceaeActinidiaceae Actinota

ceae Amentotaxaceae Adenogrammaceae Adoxaceae Aegialitidaceae Aegicerataceae

Aegilopaceae Aeginetiaceaee Aegiphilaceae Aesculaceae Aextoxicaceae Agapanthac

eae Agavaceae AgdestidaceaeAgialidaceae Agrimoniaceae Agrostidaceae Ailanthace

ae Aitoniaceae Aizoaceae Akaniaceae Alangiaceae AlchemillaceaeAldrovandaceae A

lismataceae Alliaceae Allioniaceae Allophylaceae Aloaceae Alopecuraceae Alpiniacea

e AlseuosmiaceaeAlsinaceae Alsodeiaceae Alstroemeriaceae Altingiaceae Alzateacea

e Amaranthaceae Amaryllidaceae Amborellaceae Ambrosiaceae Amentotaxaceae A

mmanniaceae Ammiaceae Amomaceae Ampelidopsaceae Amygdalaceae Amyridacea

eAnacardiaceae Anagallidaceae Anarthriaceae Anchusaceae Ancistrocladaceae Andr

omedaceae AndropogonaceaeAndrostachyaceae Androsynaceae Anemarrhenaceae

Anemonaceae Angelicaceae Anisophylleaceae AnnonaceaeAnomochloaceae Anopter

aceae Anrederaceae Anthemidaceae Anthericaceae Anthobolaceae Antidesmataceae

Antirrhinaceae Antoniaceae Aparinaceae Aphanopetalaceae Aphloiaceae Aphyllanth

aceae Apiaceae ApocynaceaeApodanthaceae Aponogetonaceae Aporusaceae Apose

ridaceae Apostasiaceae Aptandraceae AquifoliaceaeAquilariaceae Aquilegiaceae A

raceae Aragoaceae Araliaceae Aralidiaceaee Araucariaceae Arbutaceae Arceuthidace

aeArceuthobiaceae Arctostaphylaceae Arctotidaceae Ardisiaceae Arecaceae Argoph

yllaceae Arisaraceae AristolochiaceaeAristoteliaceae Arjonaceae Armeriaceae Artem

isiaceae Arthrotaxidaceae Artocarpaceae Arundinaceae ArundinellaceaeAsaraceae A

sclepiadaceae Ascyraceaee Asiraceae Asparagaceae Asperulaceae Asphodelaceae As

pidistraceaeeAsteliaceae Asteraceae Asteranthaceae Asteropeiaceae Astragalaceae

Astrocarpaceae AthanasiaceaeAtherospermataceae Atriplicaceae Atropaceae Aucuba

ceae Aurantiaceae Austrobaileyaceae Austrotaxaceae AvenaceaeAverrhoaceae Avet

raceae Avicenniaceae Azaleaceae Azimaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Balanitaceae Balanopaceae Balanophoraceae Balsameaceae Balsaminaceae Bambusa

ceae Barbaceniaceae BarbeuiaceaeBarbeyaceae Barclayaceae Barringtoniaceae Base

llaceae Bataceae Baueraceae Bauhiniaceae Baxteriaceae BegoniaceaeBehniaceae B

elangeraceae Belloniaceae Bembiciaceae Berberidaceae Berberidopsidaceae Berryac

eaee BersamaceaeBertyaceae Berzeliaceae Besleriaceae Betaceae Betulaceae Biebe

rsteiniaceae Bifariaceae Bignoniaceae BischofiaceaeBixaceae Blakeaceae Blandfordia

ceae Blattiaceae Blepharocaryaceae Blitaceae Blyxaceae Boerlagellaceae Bolivaracea

eBombacaceae Bonnetiaceae Bontiaceae Boopidaceae Boraginaceae Borassaceae B

oroniaceae BoryaceaeBotryodendraceae Bougainvilleaceae Boweniaceae Brassicacea

e Brachycaulaceae Bretschneideraceae BrexiaceaeBromeliaceae Brownlowiaceae Br

unelliaceae Bruniaceae Brunoniaceae Brunsvigiaceae Bryoniaceae BuchneraceaeBuci

daceae Buddlejaceae Buglossaceae Bulbocodiaceae Bumeliaceae Bupleuraceae Burc

hardiaceae BurmanniaceaeBurseraceae Butneriaceae Butomaceae Buxaceae Byblida

ceae Byttneriaceae

Cabombaceae Cacaoaceae Cactaceae Caesalpiniaceae Caladiaceae Calamaceae Calce

olariaceae CalectasiaceaeCalendulaceae Callaceae Callicomaceae Calligonaceae Calli

traceae Callitrichaceae Calochortaceae CalophyllaceaeCalthaceae Calycanthaceae Ca

lyceraceae Cambogiaceae Camelliaceae Campanulaceae CampynemataceaeCanacom

yricaceae Canellaceae Cannabaceae Cannaceae Canopodaceae Canotiaceae Cansjera

ceae CapparaceaeCaprariaceae Caprifoliaceae Cardiopteridaceae Carduaceae Carica

ceae Carissaceae Carlemanniaceae CarpinaceaeCarpodetaceae Cartonemataceae Ca

ryocaraceae Caryophyllaceae Casearieaceae Cassiaceae CassipoureaceaeCassythacea

e Castaneaceae Castelaceae Casuarinaceae Catesbaeaceae Cathedraceae Cecropiace

ae CedraceaeCedrelaceae Celastraceae Celosiaceae Celtidaceae Centaureaceae Cen

trolepidaceae Centroplacaceae CepaceaeCephalanthaceae Cephalotaceae Cephalota

xaceae Cerastiaceae Ceratoniaceae Ceratophyllaceae CerberaceaeCercidiphyllaceae

Cercocarpaceae Cercodiaceae Cereaceae Cerinthaceae Ceroxylaceae Cestraceae Cev

alliaceaeChailletiaceae Chamaedoraceae Chamaemoraceae Chamelauciaceae Chaun

ochitonaceae Chelidoniaceae ChelonaceaeChenopodiaceae Chimonanthaceae Chingi

thamnaceae Chionographidaceae Chiranthodendraceae ChironiaceaeChlaenaceae Ch

loanthaceae Chloranthaceae Chloridaceae Chlorogalaceae Chrysobalanaceae Cicerac

eae CichoriaceaeCimicifugaceae Cinchoniaceae Circaeaceae Circaeasteraceae Cissac

eae Cistaceae Citraceae Clematidaceae CleomaceaeClethraceae Clusiaceae Cneorac

eae Cnestidaceae Cnicaceae Cobaeaceae Cochlospermaceae Cocosaceae Coffeaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Colchicaceae Coleogynaceae Columelliaceae Combretaceae Commelinaceae Comocl

adiaceae Compositae CompsoaceaeConantheraceae Connaraceae Conostylidaceae C

onvallariaceae Convolvulaceae Cordiaceae CoreopsidaceaeCoriandraceae Coriariacea

e Coridaceae Corispermaceae Cornaceae Corokiaceae Coronillaceae Correaceae Cor

rigiolaceaeCorsiaceae Corydalaceae Corylaceae Corynocarpaceae Coryphaceae Cost

aceae Cotyledonaceae CoulaceaeCoutariaceae Coutoubiaceae Crassulaceae Crescen

tiaceae Cressaceae Crinaceae Crocaceae CroomiaceaeCrossosomataceae Crotonacea

e Cruciferae Cryptaceae Crypteroniaceae Cryptocorynaceae CryptomeriaceaeCtenol

ophonaceae Cucurbitaceae Cunninghamiaceae Cunoniaceae Cupressaceae Curcumac

eae Curtisiaceae CuscutaceaeCyananthaceae Cyanastraceae Cynanchaceae Cyanella

ceae Cycadaceae Cyclanthaceae CyclantheraceaeCyclocheilaceae Cydoniaceae Cymo

doceaceae Cynaraceae Cynocrambaceae Cynomoriaceae Cyperaceae CyphiaceaeCyp

hocarpaceae Cypripediaceae Cyrillaceae Cyrtandraceae Cyrtanthaceae Cytinaceae

Dacrycarpaceae Dactylanthaceae Damasoniaceae Daphnaceae Daphniphyllaceae Das

ypogonaceae DatiscaceaeDaturaceae Daucaceae Davidiaceae Davidsoniaceae Decais

neaceae Deeringiaceae Degeneriaceae DelphiniaceaeDendrophthoaceae Columelliac

eae Detariaceae Dialypetalanthaceae Dianellaceae Dianthaceae DiapensiaceaeDicha

petalaceae Dichondraceae Diclidantheraceae Dicrastylidiaceae Dictamnaceae Didiere

aceae DidymelaceaeDidymocarpaceae Diegodendraceae Diervillaceae Digitalidaceae

e Dilatridaceae Dilleniaceae Dioaceae DionaeaceaeDioncophyllaceae Dioscoreaceae

Diosmaceae Diospyraceae Dipentodontaceae Diphylleiaceae Diplolaenaceae Dipsaca

ceaeDipterocarpaceae Dirachmaceae Disanthaceae Diselmaceae Dodonaeaceae Do

mbeyaceae Donatiaceae DorsteniaceaeDortmannaceae Doryanthaceae Drabaceae D

racaenaceae Dracontiaceae Drimyidaceae Droseraceae DrosophyllaceaeDryadaceae

Duabangaceae Duckeodendraceae Dulongiaceae Durantaceae Durionaceae Dysphani

aceae

Ebenaceae Eccremidaceae Ecdeiocoleaceae Echiaceae Echinopaceae Ehretiaceae Ela

eagnaceae ElaeocarpaceaeElegiaceae Elatinaceae Ellisiaceae Ellisiophyllaceae Elodea

ceae Elytranthaceae Embeliaceae Emblingiaceae EmmotaceaeEmpetraceae Encepha

lartaceae Engelhardtiaceae Enhalaceae Epacridaceae Ephedraceae Ephemeranthacea

eEpilobiaceae Epimediaceae Eragrostidaceae Eremolepidaceae Eremosynaceae Erica

ceae Erinaceae EriocaulaceaeEriogonaceae Eriospermaceae Erodiaceae Erycibaceae

Eryngiaceae Erysimaceae Erythroniaceae ErythropalaceaeErythrospermaceae Erythro

xylaceae Escalloniaceae Eschscholziaceae Eucomidaceae Eucommiaceae Eucryphiace

aeEuonymaceae Eupatoriaceae Euphorbiaceae Euphrasiaceae Euphroniaceae Eupo

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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matiaceae Eupteleaceae EuryalaceaeEustrephaceae Euthemidaceae Eutocaceae Exb

ucklandiaceae Exocarpaceae

Fabaceae Fagaceae Falcatifoliaceae Ferulaceae Festucaceae Ficaceae Ficoidaceae Fit

zroyaceae FlacourtiaceaeFlagellariaceae Flindersiaceae Foetidiaceae Forestieraceae

Fothergillaceae Fouquieriaceae Fragariaceae FrancoaceaeFrangulaceae Frankeniacea

e Fraxinaceae Fraxinellaceae Freycinetiaceae Fritillariaceae Fuchsiaceae Fumariacea

eFunkiaceae

Gaiadendraceae Galacaceae Galanthaceae Galaxiaceae Galedupaceae Galeniaceae G

aliaceae Garciniaceae GardeniaceaeGardneriaceae Garryaceae Geissolomataceae Ge

itonoplesiaceae Gelsemiaceae Geniostomaceae GentianaceaeGeonomataceae Geosir

idaceae Geraniaceae Gerrardinaceae Gesneriaceae Gethyllidaceae Gilliesiaceae Gina

lloaceaeGinkgoaceae Gisekiaceae Gladiolaceae Glaucidiaceae Glechomaceae Glinace

ae Globulariaceae Gnaphaliaceae GnetaceaeGoetzeaceae Gomortegaceae Gomphiac

eae Gomphrenaceae Gonystylaceae Goodeniaceae Gordoniaceae GouaniaceaeGoup

iaceae Gramineae Gratiolaceae Grewiaceae Greyiaceae Grielaceae Griseliniaceae Gr

onoviaceae GrossulariaceaeGrubbiaceae Guaiacanaceae Guamatelaceae Guettardac

eae Gunneraceae Gustaviaceae Guttiferae GyrocarpaceaeGyrostemonaceae

Hachetteaceae Haemanthaceae Haemodoraceae Halesiaceae Hallieraceae Halocarpa

ceae Halophilaceae HalophytaceaeHaloragaceae Hamamelidaceae Hanguanaceae H

aptanthaceae Harmandaceae Hebenstretiaceae HectorellaceaeHederaceae Hedyosm

aceae Hedyotidaceae Hedysaraceae Heisteriaceae Heleniaceae Heliamphoraceae H

elianthaceaeHelianthemaceae Helichrysaceae Heliconiaceae Helicteraceae Heliotropi

aceae Helleboraceae Heloniadaceae HelosidaceaeHelwingiaceae Hemerocallidaceae

Hemimeridaceae Hemiphylacaceae Henriqueziaceae Henslowiaceae Hermanniaceae

Hernandiaceae Herniariaceae Herreriaceae Hesperocallidaceae Heterantheraceae He

teropyxidaceae HeterostylaceaeHibbertiaceae Hibiscaceae Hilleriaceae Himantandra

ceae Hippocastanaceae Hippocrateaceae HippomanaceaeHippophaeaceae Hippurida

ceae Hirtellaceae Holacanthaceae Homaliaceae Hopkinsiaceae Hoplestigmataceae H

ordeaceaeHornschuchiaceae Hortensiaceae Hortoniaceae Hostaceae Houstoniaceae

Huaceae Huerteaceae HugoniaceaeHumbertiaceae Humiriaceae Hyacinthaceae Hyd

atellaceae Hydnoraceae Hydrangeaceae Hydrastidaceae HydrillaceaeHydrocharitacea

e Hydrocotylaceae Hydrogetonaceae Hydroleaceae Hydropeltidaceae Hydrophylacac

eae HydrophyllaceaeHydrostachyaceae Hymenocardiaceae Hypecoaceae Hyperanth

eraceae Hypericaceae Hypopityaceae HypoxidaceaeHypseocharitaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta

ceae ImperatoriaceaeInocarpaceae Inulaceae Ionidiaceae Iriarteaceae Iridaceae Irvi

ngiaceae Isnardiaceae Isophysidaceae IteaceaeStrasburgeriaceae Ixiaceae Ixioliriace

ae Ixonanthaceae

Jalapaceae Jamboliferaceae Japonoliriaceae Jasionaceae Jasminiaceae Johnsoniaceae

Joinvilleaceae JuglandaceaeJulianiaceae Juncaceae Juncaginaceae Juniperaceae Jus

siaeaceae Justiciaceae

Kadsuraceae Kaliphoraceae Kalmiaceae Kaniaceae Kiggelariaceae Kingdoniaceae Kin

giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae

Krameriaceae

Labiatae Lacandoniaceae Lachenaliaceae Lacistemataceae Lactoridaceae Lactucaceae

Lagerstroemiaceaee LamiaceaeLampsanaceae Lanariaceae Langsdorffiaceae Lantan

aceae Lapageriaceae Lardizabalaceae Lasiopetalaceae LauraceaeLawsoniaceae Laxm

anniaceae Lecythidaceae Ledaceae Ledocarpaceae Leeaceae Leguminosae Leitneriac

eae LemnaceaeLennoaceae Lentibulariaceae Lentiscaceae Leoniaceae Leonticaceae

Lepidobotryaceae LepidocarpaceaeLepidocaryaceae Lepidocerataceae Leptaulaceae

Leptospermaceae Lepturaceae Lepuropetalaceae LeuctenbergiaceaeLibocedraceae

Licaniaceae Ligustraceae Lilaeaceae Liliaceae Limeaceae Limnanthaceae Limnocharit

aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace

ae Linderniaceae Lindleyaceae Linnaeaceae LippayaceaeLiriaceae Liriodendraceae Li

ssocarpaceae Littorellaceae Loasaceae Lobeliaceaee Loganiaceae LomandraceaeLoni

ceraceae Lopeziaceae Lophiocarpaceae Lophiolaceae Lophiraceae Lophophytaceae L

ophopyxidaceae LoranthaceaeLotaceae Lowiaceae Lupulaceaee Luxemburgiaceae Lu

zuriagaceae Lychnidaceae Lyciaceae LyginiaceaeLygodisodeaceae Lysimachiaceae Ly

thraceae

Macarisiaceae Mackinlayaceae Madiaceae Maesaceae Magnoliaceae Malaceae Male

sherbiaceae MalortieaceaeMalpighiaceae Malvaceae Mangiaceae Manicariaceae Ma

paniaceae Marantaceae Marathraceae MarcgraviaceaeMartyniaceae Mastixiaceae

Matricariaceae Maundiaceae Mayacaceae Medeolaceae Medusagynaceae Peridiscac

eaeMelaleucaceae Melampyraceae Melanophyllaceae Melanthiaceae Melastomatac

eae Meliaceae Melianthaceae MelicaceaeMeliosmaceae Melittidaceae Melochiacea

e Memecylaceae Mendonciaceae Menispermaceae Menthaceae MenyanthaceaeMe

nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

Nypaceae Nyssaceae

Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

ceae Papilionaceae Pappophoraceae PapyraceaeParacryphiaceae Parasitaxaceae Par

ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

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ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 22: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

22

The Main Tree above is a conservative summary of well supported relationships

between the major clades of extant seed plants Reading - interpreting and

understanding - such a phylogenetic tree is not necessarily easy so reading a tree may

require a bit of practice OHara (eg 1988 1992) provides an accessible introduction to

thinking and talking about history in the context of looking at phylogenetic trees More

recently Krell and Cranston (2004) Crisp and Cook (2005) and others have emphasized

how careful one must be when interpreting and talking about evolution in ladderized

trees in particular and phylogenetic trees in general Gregory (2008) provides a

particularly detailed discussion of this problem In ladderized trees the smaller (in terms

of numbers of terminals) sister taxon is consistently shown on the same side at every

node thus the trees tend to be pectinate like the teeth of a comb however the

horizontal axis that is as a result so evident has no polarity or particular direction The

use of the adjective basal when discussing phylogenetic trees is especially dangerous

(see also Wojciechowski et al 2004 D Soltis et al 2005b) When I use the term and the

context is not otherwise clear I am referring to the pectinations at the base of a

ladderised tree Remember that when talking about sister taxa one can never be basal

to the other or older or younger although members of one can be more derived - in the

sense of having more apomorphies - than the other (but even then one has to be

careful) The words primitive and advanced and lower and higher should

practically never be used especially when talking about taxa Amborellaceae and

Pinaceae are sister to all other flowering plants and Pinales and are basal to the crown

groups of all other flowering plantsPinales respectively but that does not mean they

are thereby primitive Pinaceae in particular have numerous apomorphies The word

plesiomorphic is far less loaded than primitive and can be used to talk about

individual characters

Most trees here have been more or less ladderized as is the one above The

ladderization may be imperfect for example you can see that on the Main Tree

asterids with ten orders follow rosids and relatives which have seventeen although

this is in part because of the currently unresolved nature of relationships between the

main clades of core eudicots (this is likely to change soon) The Main Tree could have

been drawn with Amborella Acorus or a host of other taxa at the far right without

offending any relationships phylogenetic trees are like mobiles the only fixed points

being the nodes Despite the problems just mentioned pectination interpreted

carefully has its value and it is not an arbitrary process As one reads the terminals of a

pectinate tree from left to right adjacent terminals will be separated by apomorphies

that will cumulate in an appropriate fashion When thinking of a book or a herbarium

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

23

sequence (see Haston et al 2007) this is of some value Nymphaeales and

Austrobaileyales are here adjacent on the tree but they could be separated by

hundreds of families in the sequence merely by rotating only the node from which

Nymphaeales and their sister group (all other angiosperms minus about 75 species)

arise If adjacent in a book or herbarium or as in these summary pages then it is

relatively easy to relate their characters both apomorphic and plesiomorphic but if

separated by hundreds of pages or two floors in a large building then it is less easy to

get anything from the sequence (Since all orders in the pages of the main part of this

site are preceded by the apomorphies of all nodes immediately below them in the seed

plant phylogeny and because of the linkages that have been built in to the site this

problem is much less serious there there is no sequence) So the particular sequence of

all seed plants that is based on a phylogenetic classification but is to be used in herbaria

or any other place where taxa have to be arranged linearly as in this part of APweb

will be one that maximizes the number of taxa that are both successive branches of the

tree and placed successively in the sequence Since specimens are generally filed under

families the outline of a new family sequence for arranging herbaria and books can now

be suggested (Haston et al 2007) and such a standardized sequence will further help

teaching and learning about plants

ON THINKING ABOUT APOMORPHIES

Identifying apomorphies is important because understanding the evolution of

morphology in the broad sense - ie including anatomy chemistry etc - in the context

of diversification is one of our major goals For identifying apomorphies several

preconditions must be met One needs to have an accurate well supported phylogeny

one has to have examined the right taxa both from the point of view of morphology and

molecules one has to have coded the characters right (ie delimited states

appropriately) and one has to use an appropriate model of evolution when fitting the

variation to the tree (see eg Omland 1999 Stevens 2006b) If the distributions of

apomorphies below are compared with those in other studies (eg Turgeon et al 2001

Bremer et al 2001 Endress 2001 Albach et al 2001a Judd amp Olmstead 2004 D Soltis

et al 2005b Zhang et al 2006 Judd et al 2007) the reader may find a number of

differences Although I have integrated such studies as far as is possible there are five

reasons that link to the preconditions just mentioned why there may be differences

where features are placed on trees

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

24

1 I may not have found all the information about a particular character there may be

disagreement over its interpretation or I have added information

2 The sampling of nearly all molecular studies is very incomplete (see Salisbury amp Kim

2001 for problems caused by sampling) indeed it is commonplace to decry the

incompleteness of molecular sampling But not only is the sampling in molecular studies

often less than we might wish that of the morphological and chemical characters whose

evolution we are interested in understanding is also often very poor So for many

anatomical chemical and embryological characters that are confidently said to

characterise families and other groups we all too often have no idea if those characters

are applicable to the whole clade or just to a subgroup within that clade However as

our sampling improves we can locate changes on the tree more precisely if in

unexpected positions Thus is is sometimes suggested that Ericaceae have ellagic acid

(Soltis et al 2005b) in fact the little evidence we have indicates that only a small clade

of some 80 species has ellagic acid and the rest of the family - itself alone about one

third of all Ericales - does not Similarly Ericaceae are noted for their

ectendomycorrhizal associations but Enkianthus sister to all other Ericaceae

apparently lacks these (Abe 2005) Thus these mycorrhizae and their associated

distinctive hair roots are unlikely to be apomorphies of Ericaceae - however it appears

that other families near Ericaceae may also have ectendomycorrhizae (see Asai 1934)

3 I am fitting characters to a very conservative tree with many polytomies although the

nodes that are utilised are for the most part strongly supported polytomies make the

optimisation of characters that is the assigment of character state change to a

particular node on the tree notably difficult (eg Madison amp Madison 2002) In nearly

all studies of the evolution of characters distributions of characters are optimised on a

more or less fully resolved tree and the construction of supertrees may yield yet more

detailed hypotheses of relationships (for literature on supertrees see Cotton amp

Wilkinson 2007 2008) Of course some nodes on such fully resolved trees andor

supertrees may have little support and optimisations of characters on such trees may

carry correspondingly little conviction Even parts of some of the trees used here have

poor support eg relationships within aquatic Alismatales etc although I indicate

when this is the case

4 Exactly how one goes about optimising a character on a tree is critically important

Even using simple parsimony optimisations (ACCTRAN or DELTRAN ACCelerated

TRANsitions or DELayed TRANsitions) the position of synapomorphies on trees - and

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

25

hence our ideas of evolution - may differ and this is still more true if one uses maximum

likelihood or Bayesian methods and makes apparently reasonable suggestions about

weighting gains over losses (eg Donoghue amp Ackerley 1996 Cunningham et al 1998

Omland 1997 1999 Ree amp Donoghue 1999 Polly 2001 Webster amp Purvis 2001

Ronquist 2004 Crisp amp Cook 2005) Sannier et al (2007) show how in in Arecaceae that

where on a phylogenetic tree one might peg changes in microsporogenesis will depend

on the methods one uses to do this and Pedersen et al (2007) discuss the sometimes

very substantial effect of node support on the posterior probabilities of ancestral

character states Here I use parsimony optimization not always as explicit as it might be

but I have often indicated where there are particularly important uncertainties as to the

positions of particular character changes on the tree

5 Finally although I have paid quite a lot of attention to the delimitation of the

character states that make up all the characterizations I have not spent enough time on

this critical operation If we are interested in understanding evolution then fitting the

basic variation - not character states - to a tree in principle allows greater flexibility in

understanding morphology in the context of local phylogenies (see also Stevens 2000

Endress 2005c) However many character states used here are delimited globally that

is they are circumscribed in the context of the variation shown by individual characters

across all angiosperms andor in the context of classic ideas of character evolution

Character states often have arbitrary limits and serve best to communicate

information whether they are in fact suitable for either phylogenetic analysis or

understanding evolution are separate issues Studies have rather unsurprisingly

perhaps but importantly shown that dividing the one character into different sets of

states may yield differing ideas of evolution of that character (eg Lamb Frye amp Kron

2003 Hibbett 2004) When looking at trees on which character states are optimised

one should bear in mind the problems surrounding the delimitation of states (eg

Stevens 2000 2006b) and the danger of using pollen or other types - constructs

based on many characters that vary independently but which effectively get lost in

these types thus Blackmore et al (2009) decomposed the pollen types in Asteraceae

into 52 characters

I use Remanes three main criteria of homology or better similarity when

determining the basic similarity of structures on different organisms (see Remane 1952)

These criteria are special properties position and intermediates Special properties

include anatomical or chemical characters gene expression data etc Position refers

to the position of an organ with respect to landmarks on the plant Although plants are

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

26

plants and landmarks are not as fixed as one might like trying to work out the relative

positions of parts is a good way of understanding morphology so long as one does not

try to out-do Procrustes Intermediates include intermediates found on the same

plant or on different plants Intermediates may observed during development or be

adult structures Thus one may find changes during development which suggest what

the nature of a particular structure is and very different stuctures on different plants

may turn out to be similar early in development Or one can simply compare different

taxa and see that structures that appear to be very different when comparing two

different taxa can be linked morphologically by looking at other taxa

Nevertheless the use of these criteria may not yield an unambiguous answer as to

what a structure is even given a solid phylogeny and an improved understanding of

development (see Jaramillo amp Kramer 2007 for a useful discussion) As Endress (2005c)

observed a number of features - position function development shape anatomy

histology gene activity and relationships to other taxa that clearly have petals - can be

used to distinguish a petal (for example) from other floral structures if a petal does not

have one of these features is it thereby not a petal Thus Maturen et al (2005) recently

found that floral organ diversity genes (B and C) were expressed in the large white

inflorescence bracts of Cornus (see also Costa et al 2005) Peney et al (2005) noted that

not all monosulcate pollen grains in monocots have the same developmental pathway

and that as a result such pollen might not have the same ancestral state Reeves and

Olmstead (2003) suggested that the genetic mechanisms causing monosymmetry in

Lamiales and Solanales were different and Serna and Martin (2006) described similar

problems with the development of hairs in Arabidopsis when compared with that of

hairs in Antirrhinum and Solanaceae Indeed as one perhaps might expect delimitation

of states and characters does not necessarily become easier with increasing knowledge

of development etc Thus Buzgo et al (2004) Matthews and Endress (2005) and others

have shown how hard it can be to distinguish between eg prophylls and other floral

structures as their behaviour is studied during the course of the development of the

flower Similarly at what concentration is a particular secondary metabolite deemed to

be present (Waterman (2007)

The validity of the approach used here that of fitting morphological variation to a

largely molecular-based tree may be questioned However I think it rather unlikely that

well-supported molecular branches will be overturned by morphological data Indeed

analyses of morphological data alone do provide support for many of the clades evident

in molecule-only analyses and analyses with morphological and molecular data

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

27

together may lead to increased support for clades (eg Hufford 1992 Nandi et al 1998

[but adding morphological data reduces support for a number of critical clades too]

Doyle amp Endress 2000 however in none of these papers is the use of morphology

without ambiguity) It is unfortunately clear that the use of morphology alone may not

suggest problems in the phylogenetic placement of taxa that later turn out to have been

wrongly included (eg Zhang et al 1992) Thus I have been wary of putting much weight

on clades that have only morphological support but note that I have treated molecule-

based clades with low bootstrap or jacknife support values (esp below 70) or low

posterior probabilies (below 095) likewise Although I may have been mistaken in

placing so much emphasis on molecular data in terms of providing the basic

phylogenetic framework for angiosperms morphological and molecular data are only

very rarely in irreconcilably strong conflict There are indeed a few places where the

conflict seems extreme These include the relative positions of the Monimiaceae and

Hernandiaceae (Laurales) the position of Hanguanaceae (Commelinales [as here] or

Zingiberales) and of Triplostegia (is it in Dipsacaceae or Valerianaceae - see

Dipsacales) Fossils are unlikely to affect the topologies of the trees presented here but

see below for their importance in understanding morphological evolution in general and

the evolution of angiosperms in particular Nevertheless some largely reject the idea

that trees based on molecular data alone can recover phylogenetic relationships

especially when branching points are old and prefer to used trees based on analysis of

morphological data including those taken from fossils (Hilton amp Bateman 2006 Farjon

2007)

All in all however the extent of the congruence between morphological and

molecular data is impressive and heartening and many clades can be characterised

morphologically It seemed in 1998 that there were no unambiguous morphological

synapomorphies for angiosperm orders (K Bremer 2000) and this is still true if by

unambiguous is meant non-homoplasious However many orders and other clades

have synapomorphies even if these may be indistinguishable at least at the current

level of morphological and developmental knowledge from parallel occurences

elsewhere As our morphological knowledge increases so too does the number of

apomorphies

Endress and Matthews (2006a) emphasize the importance of tendencies and

developmental constraints when thinking about characters of clades within the rosids -

many characters or character combinations occur in a rather sporadic fashion within

the clade and apparently notably less frequently outside it There are a number of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

28

examples of tendencies here and in the individual order pages such as the features

enclosed by parentheses in groups above the level of family eg see (cuticular waxes

as aggregated rodlets) for the commelinids and the discussion of the distribution of

polyandry that is flowers with numerous stamens in the asterid I + II groups etc

Tendencies may involve more than single characters When thinking of the

associations of features like integument thickness and vascularization of the integument

(Endress amp Matthews 2006a) one can imagine fairly simple developmental

preconditions being operative As Endress and Matthews (2006a) note it is difficult to

imagine vascular tissue developing in an integument only two cells thick Changes here

would be loosely correlated if morphologically linked However the strongly correlated

changes noted by Givnish et al (2005) are ecologically linked but are presumably

morphologicallydevelopmentally independent When there seem to be characters

evolving more or less together tests can be carried out to see if the changes are

concentrated on certain branches of the tree (eg Maddison 1990 Sanderson 1991

Maddison amp Maddison 2000) Such tests have rarely been carried out at higher levels in

angiosperms

One of the most striking examples of a tendency is the distribution of N-fixation

restricted as it is to a monophyletic group of four clades although it has arisen seven (or

perhaps several more) times independently within the N-fixing clade and several

members of two quite different kinds of bacteria are involved (eg Clawson et al 2004

Elliott et al 2007 Sprent amp James 2007) Other examples of work that bears on the issue

of tendencies include the findings that flowers of polysymmetrical Arabidopsis have

genes like TCP1 that are expressed asymmetrically during early development and TCP1

is a probable orthologue of the well-known CYC gene of Antirrhinum that is involved in

the development of monosymmetric slowers there (Cubas et al 2001 Costa et al 2005

etc) Parallelism might build on this underlying morphologically cryptic monosymmetry

even if details of the genetic mechanisms causing the monosymmetry evident in

particular groups may be different (Reeves amp Olmstead 2003 Cubas 2004 see above)

Similarly there have been several recent suggestions that the capability to synthesise a

particular metabolite may be switched off but not lost and so can sometimes be

reacquired (eg Wink amp Witte 1983 Wink 2003 Liscombe et al 2005 Larsson 2007

Waterman 2007) Hence perhaps the rather spotty distribution of many secondary

metabolites like ellagic acid the indole alkaloid camptothecin iridoids etc when

considered in the context of phylogenies As a non-botanical example - but a rather nice

one - Salwini-Plawen and Mayr (1961) suggested some time ago that there has been

considerable parallelism (40-65 or more independent origins) in the evolution of eyes in

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

29

metazoans However Pax 6 seems to be a master control gene that is involved in eye

formation perhaps throughout all bilateralians (eg Gehring amp Ikeo 1999 Erwin amp

Davidson 2002) which changes the nature of the problem Note however that in

general our invocation of evolutionary tendencies to explain the patterns of

distributions of characters is really an admission that we do not understand those

patterns

As we find out more about variation we will probably find fewer and fewer features

constant throughout a group Most unqualified statements of presence and absence

should properly be qualified as usually present or usually absent if one is thinking of

the characterisations on the order pages as encompassing the total variation within a

clade Thus Pistia alone among monocots as so far known has sieve tube plastids with

starch grains not protein crystals However this does not affect the fact that sieve tube

plastids with cuneate protein crystals and lacking starch grains are an apomorphy for

monocots Furthermore fossils in a number of cases suggest character combinations

unknown in extant taxa as may be seen in the discussions of Fagaceae Platanaceae

Iteaceae Calycanthaceae etc Confusing the issue there may be questions as to where

exactly on the tree a particular fossil is to be placed (eg see Nymphaeaceae

Calycanthaceae Archaefructus etc)

To summarize given our current understandings of both phylogenies and

characters evolution of some characters in which we are interested seems very labile

(see eg D Soltis et al 2005b Endress and Matthews 2006a Stevens 2006b Ekman et

al 2008 for a good recent study) and I have been cautious when talking about character

evolution Much effort must continue to be spent in summarizing characters of well-

established clades at all levels providing features by which they may be recognized and

signaling synapomorphies Remember that (1) the basic morphological anatomical and

chemical knowledge of many critical taxa is woefully incomplete (2) different

assumptions about character evolution may greatly affect the position of

synapomorphies on trees (3) in many cases relationships within and between many

groups are too uncertain at present to worry very much about synapomorphies and

(4) we must be clear about what we do and do not not know As mentioned above I

have indicated in a number of places where there are particularly important

uncertainties as to where characters should be placed on the tree Nevertheless it is a

relatively easy matter to update notes such as these and it can be a simple matter to

incorporate new data on characters that have never before been considered in the

context of a tree Much basic - and unfortunately perhaps unfashionable - work must be

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

30

carried to clarify the distribution of morphological anatomical and chemical characters

for examples of what can to be done see the work of P K Endress and collaborators

(floral morphology and development) and S R Jensen and collaborators (iridoids) But

acquiring information about nodal anatomy cell and tissue distribution nodal anatomy

and the like is not difficult at all and should be generally encouraged

But all this is merely a necesary prelude to the understanding of evolution For this

one needs to know a lot more including dating the phylogeny understanding the

function(s) of characters factoring in relevant aspects of the palaeoenvironment in

which an apomorphy first appeared etc

SUMMARY OF THE SYSTEM

Below is a formalised summary of the relationships within orders of the families of seed

plants There are a few families that are not recognised even as options in APG II and

vice versa as well as a few extra orders The families to be recognised in the most

recent edition of Mabberleys The Plant Book (Mabberley 2008) are also largely

consistent with those below But all differences are trivial and will - I hope - eventually

disappear

Square brackets - [] - enclose clades the plus sign - + - designates sister taxa a

comma - - denotes part of a polytomy and quotation marks - - denotes a

paraphyletic group

SEED PLANTS

GYMNOSPERMS

Cycadales

Cycadaceae + Zamiaceae

Ginkgoales

Ginkgoaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

31

Pinales

Pinaceae [[Araucariaceae [Podocarpaceae + Phyllocladaceae]] [Sciadopityaceae

[Taxaceae + Cupressaceae]]]

Gnetales

Ephedraceae [Gnetaceae + Welwitschiaceae]

ANGIOSPERMSFLOWERING PLANTS

Amborellales

Amborellaceae

Nymphaeales

Hydatellaceae [Cabombaceae + Nymphaeaceae]

Austrobaileyales

Austrobaileyaceae [Schisandraceae + Trimeniaceae]

Chloranthales

Chloranthaceae

MAGNOLIIDS

Magnoliales

Myristicaceae [Magnoliaceae [[Himantandraceae + Degeneriaceae] [Eupomatiaceae +

Annonaceae]]]

Laurales

Calycanthaceae [[Siparunaceae [Gomortegaceae + Atherospermataceae]] [Monimiaceae

[Hernandiaceae + Lauraceae]]]

Canellales

Canellaceae + Winteraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

32

Piperales

[Hydnoraceae + Aristolochiaceae] [Piperaceae + Saururaceae]

MONOCOTS

Acorales

Acoraceae

Alismatales

Araceae [Tofieldiaceae [[Alismataceae [Hydrocharitaceae + Butomaceae]]

[Scheuchzeriaceae [Aponogetonaceae [Juncaginaceae [Maundiaceae [[Posidoniaceae

[Ruppiaceae + Cymodoceaceae]] [Zosteraceae + Potamogetonaceae]]]]]]]

Petrosaviales

Petrosaviaceae

Dioscoreales

Nartheciaceae [[Taccaceae + Thismiaceae] [Burmanniaceae + Dioscoreaceae]]

Pandanales

Velloziaceae Triuridaceae Stemonaceae [Pandanaceae + Cyclanthaceae]

Liliales

Corsiaceae [Campynemataceae [Petermanniaceae [Colchicaceae + Alstroemeriaceae]]

Melanthiaceae [[Philesiaceae + Rhipogonaceae] [Smilacaceae + Liliaceae]]]

Asparagales

Orchidaceae [[Boryaceae [Blandfordiaceae [Lanariaceae [Asteliaceae + Hypoxidaceae]]]]

[[Ixioliriaceae + Tecophilaeaceae] [Doryanthaceae [Iridaceae [Xeronemataceae

[Xanthorrhoeaceae [Alliaceae + Asparagaceae]]]]]]]

COMMELINIDS

Unplaced

Dasypogonaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

33

Arecales

Arecaceae

Poales

[Typhaceae + Bromeliaceae] [Rapateaceae [[Thurniaceae [Juncaceae + Cyperaceae]]

[[Mayacaceae [Eriocaulaceae + Xyridaceae]] [Flagellariaceae [Anarthriaceae

[Centrolepidaceae + Restionaceae]]] [Joinvilleaceae [Ecdeiocoleaceae + Poaceae]]]]]]]

Commelinales

[Commelinaceae + Hanguanaceae] [Philydraceae [Haemodoraceae + Pontederiaceae]]

Zingiberales

Musaceae [Strelitziaceae + Lowiaceae] Heliconiaceae [[Cannaceae + Marantaceae]

[Costaceae + Zingiberaceae]]

Ceratophyllales

Ceratophyllaceae

EUDICOTS

Ranunculales

Eupteleaceae [Papaveraceae [[[Lardizabalaceae + Circaeasteraceae] [Menispermaceae

[Berberidaceae + Ranunculaceae]]]]

Sabiales

Sabiaceae

Proteales

Nelumbonaceae [Platanaceae + Proteaceae]

Trochodendrales

Trochodendraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

34

Buxales

Haptanthaceae [Buxaceae + Didymelaceae]

CORE EUDICOTS

Gunnerales

Gunneraceae + Myrothamnaceae

Dilleniales

Dilleniaceae

Saxifragales

Peridiscaceae [[Paeoniaceae [Altingiaceae [Hamamelidaceae [Cercidiphyllaceae +

Daphniphyllaceae]]]] [[Crassulaceae [Aphanopetalaceae [Tetracarpaeaceae

[Penthoraceae + Haloragaceae]]]] [Iteaceae [Grossulariaceae + Saxifragaceae]]]]

Cynomoriaceae unplaced

Vitales

Vitaceae

ROSIDS

FABIDROSID I

Zygophyllales

Krameriaceae + Zygophyllaceae

Celastrales

Lepidobotryaceae + Celastraceae

Oxalidales

Huaceae [[Connaraceae + Oxalidaceae] [Cunoniaceae [Elaeocarpaceae [Brunelliaceae +

Cephalotaceae]]]]

Malpighiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

35

[Achariaceae [Goupiaceae [Violaceae + Passifloraceae] [Lacistemataceae Salicaceae]]

[Lophopyxidaceae Putranjivaceae] [Ctenolophonaceae [Erythroxylaceae +

Rhizophoraceae]] Linaceae Ixonanthaceae Humiriaceae Irvingiaceae

Centroplacaceae Pandaceae [[Bonnetiaceae + Clusiaceae] [Calophyllaceae

[Hypericaceae + Podostemaceae]]] [Malpighiaceae + Elatinaceae] Ochnaceae

[Peraceae [Rafflesiaceae + Euphorbiaceae]] [Phyllanthaceae + Picrodendraceae]

[Balanopaceae [[Trigoniaceae + Dichapetalaceae] [Chrysobalanaceae +

Euphroniaceae]]] Caryocaraceae

N-FIXING CLADE

Fabales

Quillajaceae [Fabaceae [Polygalaceae + Surianaceae]]

Rosales

Rosaceae [[Barbeyaceae [Elaeagnaceae Dirachmaceae Rhamnaceae]] [Ulmaceae

[Cannabaceae [Moraceae + Urticaceae]]]]

Cucurbitales

Anisophylleaceae [[Corynocarpaceae + Coriariaceae] [Cucurbitaceae [Tetramelaceae

[Datiscaceae + Begoniaceae]]]] Apodanthaceae

Fagales

Nothofagaceae [Fagaceae [[Myricaceae + Juglandaceae] [Casuarinaceae

[Ticodendraceae + Betulaceae]]]]

MALVIDROSID II

pgtGeraniales

Geraniaceae [[Melianthaceae + Francoaceae] [Vivianaceae + Ledocarpaceae]]

Myrtales

Combretaceae [[Onagraceae + Lythraceae] [[Vochysiaceae + Myrtaceae]

[Melastomataceae [Crypteroniaceae [Alzateaceae + Penaeaceae]]]]]

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

36

Crossosomatales

[Staphyleaceae [Guamatelaceae [Crossosomataceae + Stachyuraceae]]] [Aphloiaceae

[Geissolomataceae + Strasburgeriaceae]]

Picramniales

Picramniaceae

Sapindales

Biebersteiniaceae Nitrariaceae [Kirkiaceae [Anacardiaceae + Burseraceae]]

[Sapindaceae [Simaroubaceae Rutaceae Meliaceae]]

Huerteales

Gerradinaceae [Dipentodontaceae + Tapisciaceae]

Malvales

Neuradaceae [Thymelaeaceae [Sphaerosepalaceae Bixaceae [Cistaceae

[Sarcolaenaceae + Dipterocarpaceae]] [Cytinaceae + Muntingiaceae] Malvaceae]]

Brassicales

[Akaniaceae + Tropaeolaceae] [[Moringaceae + Caricaceae] [Setchellanthaceae

[Limnanthaceae [[Koeberliniaceae [Bataceae + Salvadoraceae]] [Emblingiaceae

[Pentadiplandraceae [Gyrostemonaceae Resedaceae Stixis and relatives] Tovariaceae

[Capparaceae [Cleomaceae + Brassicaceae]]]]]]]]

Berberidopsidales

Aextoxicaceae + Berberidopsidaceae

Santalales

Erythropalaceae Olacaceae [[Loranthaceae [Misodendraceae + Schoepfiaceae]]

[Opiliaceae + Santalaceae]] Balanophoraceae unplaced

Caryophyllales

[[Droseraceae [Nepenthaceae [Drosophyllaceae [Ancistrocladaceae +

Dioncophylleaceae]]]]] [[Frankeniaceae + Tamaricaceae] [Polygonaceae +

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

37

Plumbaginaceae]]] [Rhabdodendraceae [Simmondsiaceae [[Asteropeiaceae +

Physenaceae] [[Caryophyllaceae [Achatocarpaceae + Amaranthaceae]]

[Stegnospermataceae [Limeaceae [[Lophocarpaceae [Barbeuiaceae [Aizoaceae

[Sarcobataceae Phytolaccaceae Nyctaginaceae]]]] [Molluginaceae [Halophytaceae

Montiaceae Basellaceae Didiereaceae [Talinaceae [Portulacaceae [Anacampseros etc +

Cactaceae]]]]]]]]]]]]]

ASTERIDS

Cornales

[Cornaceae + Nyssaceae] [Hydrangeaceae + Loasaceae] Hydrostachyaceae [Curtisiaceae

+ Grubbiaceae]

Ericales

[Balsaminaceae [Marcgraviaceae + Tetrameristaceae]] [[Polemoniaceae +

Fouquieraceae] Lecythidaceae [[Sladeniaceae + Pentaphylacaceae] [Sapotaceae

[Ebenaceae [Maesaceae [Theophrastaceae [Myrsinaceae + Primulaceae]]]]]

[Mitrastemonaceae Theaceae [Symplocaceae [Styracaceae + Diapensiaceae]]

[[Sarraceniaceae [Roridulaceae + Actinidiaceae] [Clethraceae [Cyrillaceae +

Ericaceae]]]]]]

LAMIIDASTERID I

Unplaced

Metteniusaceae Oncothecaceae Icacinaceae and some unplaced ex-Icacinaceae

Garryales

Garryaceae + Eucommiaceae

Unplaced

Boraginaceae Vahliaceae

Gentianales

Rubiaceae [Gentianaceae [Loganiaceae [Gelsemiaceae + Apocynaceae]]]

Lamiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

38

Plocospermataceae [Oleaceae [Tetrachondraceae [Calceolariaceae

[Gesneriaceae Peltanthera [Pedaliaceae Carlemanniaceae Martyniaceae Stilbaceae

Plantaginaceae Linderniaceae Bignoniaceae Verbenaceae Lamiaceae Paulowniaceae

Schlegeliaceae Thomandersiaceae Phrymaceae Rehmannia Acanthaceae

Scrophulariaceae Orobanchaceae Byblidaceae Lentibulariaceae]]]]]

Solanales

[Montiniaceae [Sphenocleaceae + Hydroleaceae]] [Convolvulaceae + Solanaceae]

CAMPANULIDASTERID II

Aquifoliales

[[Cardiopteridaceae + Stemonuraceae] [Aquifoliaceae [Helwingiaceae +

Phyllonomaceae]]

Asterales

[Rousseaceae + Campanulaceae] Pentaphragmataceae [[Alseuosmiaceae [Phellinaceae

+ Argophyllaceae]] Stylidiaceae [Menyanthaceae [Goodeniaceae [Calyceraceae +

Asteraceae]]]]

Escalloniales

Escalloniaceae

Bruniales

[Bruniaceae + Columelliaceae]

Apiales

Pennantiaceae [Torricelliaceae [Griseliniaceae [Pittosporaceae [Araliaceae

[Myodocarpaceae + Apiaceae]]]]]

Paracryphiales

Paracryphiaceae

Dipsacales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

39

Adoxaceae [Diervillaceae [Caprifoliaceae [Linnaeaceae [Morinaceae [Dipsacaceae +

Valerianaceae]]]]]

LINKS TO ORDERS AND FAMILIES

[Back to Top]

MAIN GROUPINGS

Asterids commelinids core eudicots asterid 1 asterid

2 eudicots gymnosperms Magnoliophyta monocots N-fixing claderosids seed plants

ALPHABETICAL LISTING OF ALL ORDINAL NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | L | M | N | O | P | Q | R | S | T | U | V | W | X | Z

Abietales Acanthales Acerales Acorales Actinidiales Actinostrobales Adoxales Aescul

ales Agavales Aizoales AkanialesAlismatales Alliales Alseuosmiales Alstroemeriales

Altingiales Amaranthales Amaryllidales Amborellales AmbrosialesAmmiales Amomal

es Ancistrocladales Anisophylleales Annonales Anthobolales Apiales Apocynales Apo

nogetonalesAquifoliales Arales Araliales Aralidiales Araucariales Arecales Aristolochi

ales Athrotaxidales Asarales AsclepiadalesAsparagales Asphodelales Asparagales Ast

eliales Atriplicales Aucubales Austrobaileyales Avenales

Balanitales Balanopales Balanophorales Balsaminales Barbeyales Barclayales Batales

Begoniales BerberidalesBerberidopsidales Betulales Biebersteiniales Bignoniales Bix

ales Boraginales Brassicales Brexiales Bromeliales BrunialesBrunoniales Burmannial

es Burserales Butomales Buxales Byblidales

Cactales Callitrichales Calycanthales Calycerales Campanulales Campynematales Can

ellales Cannales CapparalesCaprifoliales Cardiopteridales Carduales Caricales Carle

manniales Caryophyllales Cassiales Casuarinales CelastralesCentrolepidales Cephalot

ales Cephalotaxales Ceratophyllales Cercidiphyllales Chenopodiales Chironiales Chlor

anthalesChrysobalanales Cinchonales Circaeasterales Cistales Citrales Cocosales Col

chicales Columelliales CombretalesCommelinales Connarales Convolvulales Coriarial

es Cornales Corylales Corynocarpales Crassulales CrossosomatalesCucurbitales Cunn

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

40

inghamiales Cunoniales Cupressales Cycadales Cyclanthales Cymodoceales Cynarales

CynomorialesCyperales Cyrillales Cytinales

Daphnales Daphniphyllales Dasypogonales Datiscales Degeneriales Desfontainiales D

iapensiales Didymelales DillenialesDioales Dioncophyllales Dioscoreales Diospyrales

Dipentodontales Dipsacales Droserales

Ebenales Echiales Elaeagnales Elaeocarpales Elatinales Elodeales Emmotales Empetr

ales Ephedrales Ericales EriocaulalesErythropalaless Escalloniales Eucommiales Euph

orbiales Eupomatiales Eupteleales Euryalaless

Fabales Fagales Falcatifoliales Ficales Flacourtiales Flagellariales Fouquieriales Franc

oales Frangulales

Galiales Garryales Geissolomatales Gentianales Geraniales Gesneriales Ginkgoales G

laucidiales Globulariales GnetalesGoodeniales Greyiales Griseliniales Grossulariales

Grubbiales Gunnerales Gyrocarpales Gyrostemonales

Haemodorales Haloragales Hamamelidales Hanguanales Heisteriales Helleborales He

lwingiales Himantandrales HippuridalesHomaliales Hortensiales Huales Huerteales

Hydatellales Hydnorales Hydrangeales Hydrastidales HydrocharitalesHydropeltidales

Hydrostachyales Hypericales Hypoxidales

Icacinales Illiciales Iridales Irvingales Iteales Ixiales

Jasminales Juglandales Julianiales Juncaginales Juncales

Lacistematales Lactoridales Lamiales Lardizabalales Laurales Lecythidales Ledocarpal

es Leitneriales LentibularialesLigustrales Liliales Limnanthales Linales Liriales Loasal

es Lobeliales Loganiales Lonicerales Loranthales LowialesLythrales

Magnoliales Malpighiales Malvales Marathrales Marcgraviales Mayacales Medusagy

nales Medusandrales MelanthialesMelastomatales Meliales Melianthales Meliosmal

es Menispermales Menyanthales Metteniusales MitrastemonalesMiyoshiales Moni

miales Moringales Musales Myricales Myristicales Myrothamnales Myrsinales Myrta

les

Najadales Nandinales Narcissales Nartheciales Nelumbonales Nepenthales Neuradal

es Nitrariales Nolanales NothofagalesNyctaginales Nymphaeales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

41

Ochnales Oenotherales Olacales Oleales Onagrales Oncothecales Opuntiales Orchid

ales Oxalidales

Saxifragales Pandales Pandanales Papaverales Paracryphiales Parasitaxales Paridales

Parnassiales PassifloralesPenaeales Pennantiales Pentaphragmatales Peridiscales P

etiveriales Petrosaviales Phellinales Philydrales PhyllanthalesPhysenales Phytolaccal

es Picramniales Pinales Pinguiculales Piperales Pittosporales Plantaginales Platanale

sPlumbaginales Poales Podocarpales Podophyllales Podostemales Polemoniales Pol

ygalales Polygonales PontederialesPortulacales Posidoniales Potamogetonales Primu

lales Proteales

Quercales Quillajales Quintiniales

Rafflesiales Ranunculales Rapateales Resedales Restionales Rhabdodendrales Rhamn

ales Rhinanthales RhizophoralesRhodorales Rhoipteleales Roridulales Rosales Rouss

eales Rubiales Ruppiales Rutales

Sabiales Salicales Salvadorales Samolales Samydales Sanguisorbales Santalales Sapin

dales Sapotales SarracenialesSaxegotheales Saxifragales Scheuchzeriales Sciadopitya

les Scleranthales Scrophulariales Scyphostegiales SedalesSilenales Simmondsiales S

milacales Solanales Sphenocleales Sphenostemonales Stangeriales Stellariales Stemo

nalesStilbales Stylidiales Styracales Surianales

Taccales Tamales Tamaricales Taxales Taxodiales Tecophilaeales Terebinthales Tern

stroemiales Theales TheligonalesThymelaeales Tiliales Tofieldiales Torricelliales Tov

ariales Tribelales Trilliales Trimeniales Triuridales TrochodendralesTropaeolales Turn

erales Typhales

Ulmales Urticales

Vacciniales Vahliales Vallisneriales Velloziales Veratrales Verbenales Viburnales Vinc

ales Violales Viscales VitalesVochysiales

Welwitschiales Winterales

Xanthorrhoeales Ximeniales Xyridales

Zamiales Zingiberales Zosterales Zygophyllales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

42

ALPHABETICAL LISTING OF ALL FAMILY NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | K | L | M | N | O | P | Q | R | S | T | U | V | W | X | Y

|Z

Back to Home

In some cases clicking on a family synonym will send you directly to the subfamily in

which the genus of the basionym is to be found

Abietaceae Abolbodaceae Abrophyllaceae Acalyphaceae Acanthaceae [Acanthochla

mydaceae] Acarnaceaee AceraceaeAchariaceae Achatocarpaceae Achradaceae Achy

ranthaceae Acmopylaceae Acoraceae Acristaceae ActaeaceaeActinidiaceae Actinota

ceae Amentotaxaceae Adenogrammaceae Adoxaceae Aegialitidaceae Aegicerataceae

Aegilopaceae Aeginetiaceaee Aegiphilaceae Aesculaceae Aextoxicaceae Agapanthac

eae Agavaceae AgdestidaceaeAgialidaceae Agrimoniaceae Agrostidaceae Ailanthace

ae Aitoniaceae Aizoaceae Akaniaceae Alangiaceae AlchemillaceaeAldrovandaceae A

lismataceae Alliaceae Allioniaceae Allophylaceae Aloaceae Alopecuraceae Alpiniacea

e AlseuosmiaceaeAlsinaceae Alsodeiaceae Alstroemeriaceae Altingiaceae Alzateacea

e Amaranthaceae Amaryllidaceae Amborellaceae Ambrosiaceae Amentotaxaceae A

mmanniaceae Ammiaceae Amomaceae Ampelidopsaceae Amygdalaceae Amyridacea

eAnacardiaceae Anagallidaceae Anarthriaceae Anchusaceae Ancistrocladaceae Andr

omedaceae AndropogonaceaeAndrostachyaceae Androsynaceae Anemarrhenaceae

Anemonaceae Angelicaceae Anisophylleaceae AnnonaceaeAnomochloaceae Anopter

aceae Anrederaceae Anthemidaceae Anthericaceae Anthobolaceae Antidesmataceae

Antirrhinaceae Antoniaceae Aparinaceae Aphanopetalaceae Aphloiaceae Aphyllanth

aceae Apiaceae ApocynaceaeApodanthaceae Aponogetonaceae Aporusaceae Apose

ridaceae Apostasiaceae Aptandraceae AquifoliaceaeAquilariaceae Aquilegiaceae A

raceae Aragoaceae Araliaceae Aralidiaceaee Araucariaceae Arbutaceae Arceuthidace

aeArceuthobiaceae Arctostaphylaceae Arctotidaceae Ardisiaceae Arecaceae Argoph

yllaceae Arisaraceae AristolochiaceaeAristoteliaceae Arjonaceae Armeriaceae Artem

isiaceae Arthrotaxidaceae Artocarpaceae Arundinaceae ArundinellaceaeAsaraceae A

sclepiadaceae Ascyraceaee Asiraceae Asparagaceae Asperulaceae Asphodelaceae As

pidistraceaeeAsteliaceae Asteraceae Asteranthaceae Asteropeiaceae Astragalaceae

Astrocarpaceae AthanasiaceaeAtherospermataceae Atriplicaceae Atropaceae Aucuba

ceae Aurantiaceae Austrobaileyaceae Austrotaxaceae AvenaceaeAverrhoaceae Avet

raceae Avicenniaceae Azaleaceae Azimaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

43

Balanitaceae Balanopaceae Balanophoraceae Balsameaceae Balsaminaceae Bambusa

ceae Barbaceniaceae BarbeuiaceaeBarbeyaceae Barclayaceae Barringtoniaceae Base

llaceae Bataceae Baueraceae Bauhiniaceae Baxteriaceae BegoniaceaeBehniaceae B

elangeraceae Belloniaceae Bembiciaceae Berberidaceae Berberidopsidaceae Berryac

eaee BersamaceaeBertyaceae Berzeliaceae Besleriaceae Betaceae Betulaceae Biebe

rsteiniaceae Bifariaceae Bignoniaceae BischofiaceaeBixaceae Blakeaceae Blandfordia

ceae Blattiaceae Blepharocaryaceae Blitaceae Blyxaceae Boerlagellaceae Bolivaracea

eBombacaceae Bonnetiaceae Bontiaceae Boopidaceae Boraginaceae Borassaceae B

oroniaceae BoryaceaeBotryodendraceae Bougainvilleaceae Boweniaceae Brassicacea

e Brachycaulaceae Bretschneideraceae BrexiaceaeBromeliaceae Brownlowiaceae Br

unelliaceae Bruniaceae Brunoniaceae Brunsvigiaceae Bryoniaceae BuchneraceaeBuci

daceae Buddlejaceae Buglossaceae Bulbocodiaceae Bumeliaceae Bupleuraceae Burc

hardiaceae BurmanniaceaeBurseraceae Butneriaceae Butomaceae Buxaceae Byblida

ceae Byttneriaceae

Cabombaceae Cacaoaceae Cactaceae Caesalpiniaceae Caladiaceae Calamaceae Calce

olariaceae CalectasiaceaeCalendulaceae Callaceae Callicomaceae Calligonaceae Calli

traceae Callitrichaceae Calochortaceae CalophyllaceaeCalthaceae Calycanthaceae Ca

lyceraceae Cambogiaceae Camelliaceae Campanulaceae CampynemataceaeCanacom

yricaceae Canellaceae Cannabaceae Cannaceae Canopodaceae Canotiaceae Cansjera

ceae CapparaceaeCaprariaceae Caprifoliaceae Cardiopteridaceae Carduaceae Carica

ceae Carissaceae Carlemanniaceae CarpinaceaeCarpodetaceae Cartonemataceae Ca

ryocaraceae Caryophyllaceae Casearieaceae Cassiaceae CassipoureaceaeCassythacea

e Castaneaceae Castelaceae Casuarinaceae Catesbaeaceae Cathedraceae Cecropiace

ae CedraceaeCedrelaceae Celastraceae Celosiaceae Celtidaceae Centaureaceae Cen

trolepidaceae Centroplacaceae CepaceaeCephalanthaceae Cephalotaceae Cephalota

xaceae Cerastiaceae Ceratoniaceae Ceratophyllaceae CerberaceaeCercidiphyllaceae

Cercocarpaceae Cercodiaceae Cereaceae Cerinthaceae Ceroxylaceae Cestraceae Cev

alliaceaeChailletiaceae Chamaedoraceae Chamaemoraceae Chamelauciaceae Chaun

ochitonaceae Chelidoniaceae ChelonaceaeChenopodiaceae Chimonanthaceae Chingi

thamnaceae Chionographidaceae Chiranthodendraceae ChironiaceaeChlaenaceae Ch

loanthaceae Chloranthaceae Chloridaceae Chlorogalaceae Chrysobalanaceae Cicerac

eae CichoriaceaeCimicifugaceae Cinchoniaceae Circaeaceae Circaeasteraceae Cissac

eae Cistaceae Citraceae Clematidaceae CleomaceaeClethraceae Clusiaceae Cneorac

eae Cnestidaceae Cnicaceae Cobaeaceae Cochlospermaceae Cocosaceae Coffeaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

44

Colchicaceae Coleogynaceae Columelliaceae Combretaceae Commelinaceae Comocl

adiaceae Compositae CompsoaceaeConantheraceae Connaraceae Conostylidaceae C

onvallariaceae Convolvulaceae Cordiaceae CoreopsidaceaeCoriandraceae Coriariacea

e Coridaceae Corispermaceae Cornaceae Corokiaceae Coronillaceae Correaceae Cor

rigiolaceaeCorsiaceae Corydalaceae Corylaceae Corynocarpaceae Coryphaceae Cost

aceae Cotyledonaceae CoulaceaeCoutariaceae Coutoubiaceae Crassulaceae Crescen

tiaceae Cressaceae Crinaceae Crocaceae CroomiaceaeCrossosomataceae Crotonacea

e Cruciferae Cryptaceae Crypteroniaceae Cryptocorynaceae CryptomeriaceaeCtenol

ophonaceae Cucurbitaceae Cunninghamiaceae Cunoniaceae Cupressaceae Curcumac

eae Curtisiaceae CuscutaceaeCyananthaceae Cyanastraceae Cynanchaceae Cyanella

ceae Cycadaceae Cyclanthaceae CyclantheraceaeCyclocheilaceae Cydoniaceae Cymo

doceaceae Cynaraceae Cynocrambaceae Cynomoriaceae Cyperaceae CyphiaceaeCyp

hocarpaceae Cypripediaceae Cyrillaceae Cyrtandraceae Cyrtanthaceae Cytinaceae

Dacrycarpaceae Dactylanthaceae Damasoniaceae Daphnaceae Daphniphyllaceae Das

ypogonaceae DatiscaceaeDaturaceae Daucaceae Davidiaceae Davidsoniaceae Decais

neaceae Deeringiaceae Degeneriaceae DelphiniaceaeDendrophthoaceae Columelliac

eae Detariaceae Dialypetalanthaceae Dianellaceae Dianthaceae DiapensiaceaeDicha

petalaceae Dichondraceae Diclidantheraceae Dicrastylidiaceae Dictamnaceae Didiere

aceae DidymelaceaeDidymocarpaceae Diegodendraceae Diervillaceae Digitalidaceae

e Dilatridaceae Dilleniaceae Dioaceae DionaeaceaeDioncophyllaceae Dioscoreaceae

Diosmaceae Diospyraceae Dipentodontaceae Diphylleiaceae Diplolaenaceae Dipsaca

ceaeDipterocarpaceae Dirachmaceae Disanthaceae Diselmaceae Dodonaeaceae Do

mbeyaceae Donatiaceae DorsteniaceaeDortmannaceae Doryanthaceae Drabaceae D

racaenaceae Dracontiaceae Drimyidaceae Droseraceae DrosophyllaceaeDryadaceae

Duabangaceae Duckeodendraceae Dulongiaceae Durantaceae Durionaceae Dysphani

aceae

Ebenaceae Eccremidaceae Ecdeiocoleaceae Echiaceae Echinopaceae Ehretiaceae Ela

eagnaceae ElaeocarpaceaeElegiaceae Elatinaceae Ellisiaceae Ellisiophyllaceae Elodea

ceae Elytranthaceae Embeliaceae Emblingiaceae EmmotaceaeEmpetraceae Encepha

lartaceae Engelhardtiaceae Enhalaceae Epacridaceae Ephedraceae Ephemeranthacea

eEpilobiaceae Epimediaceae Eragrostidaceae Eremolepidaceae Eremosynaceae Erica

ceae Erinaceae EriocaulaceaeEriogonaceae Eriospermaceae Erodiaceae Erycibaceae

Eryngiaceae Erysimaceae Erythroniaceae ErythropalaceaeErythrospermaceae Erythro

xylaceae Escalloniaceae Eschscholziaceae Eucomidaceae Eucommiaceae Eucryphiace

aeEuonymaceae Eupatoriaceae Euphorbiaceae Euphrasiaceae Euphroniaceae Eupo

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

45

matiaceae Eupteleaceae EuryalaceaeEustrephaceae Euthemidaceae Eutocaceae Exb

ucklandiaceae Exocarpaceae

Fabaceae Fagaceae Falcatifoliaceae Ferulaceae Festucaceae Ficaceae Ficoidaceae Fit

zroyaceae FlacourtiaceaeFlagellariaceae Flindersiaceae Foetidiaceae Forestieraceae

Fothergillaceae Fouquieriaceae Fragariaceae FrancoaceaeFrangulaceae Frankeniacea

e Fraxinaceae Fraxinellaceae Freycinetiaceae Fritillariaceae Fuchsiaceae Fumariacea

eFunkiaceae

Gaiadendraceae Galacaceae Galanthaceae Galaxiaceae Galedupaceae Galeniaceae G

aliaceae Garciniaceae GardeniaceaeGardneriaceae Garryaceae Geissolomataceae Ge

itonoplesiaceae Gelsemiaceae Geniostomaceae GentianaceaeGeonomataceae Geosir

idaceae Geraniaceae Gerrardinaceae Gesneriaceae Gethyllidaceae Gilliesiaceae Gina

lloaceaeGinkgoaceae Gisekiaceae Gladiolaceae Glaucidiaceae Glechomaceae Glinace

ae Globulariaceae Gnaphaliaceae GnetaceaeGoetzeaceae Gomortegaceae Gomphiac

eae Gomphrenaceae Gonystylaceae Goodeniaceae Gordoniaceae GouaniaceaeGoup

iaceae Gramineae Gratiolaceae Grewiaceae Greyiaceae Grielaceae Griseliniaceae Gr

onoviaceae GrossulariaceaeGrubbiaceae Guaiacanaceae Guamatelaceae Guettardac

eae Gunneraceae Gustaviaceae Guttiferae GyrocarpaceaeGyrostemonaceae

Hachetteaceae Haemanthaceae Haemodoraceae Halesiaceae Hallieraceae Halocarpa

ceae Halophilaceae HalophytaceaeHaloragaceae Hamamelidaceae Hanguanaceae H

aptanthaceae Harmandaceae Hebenstretiaceae HectorellaceaeHederaceae Hedyosm

aceae Hedyotidaceae Hedysaraceae Heisteriaceae Heleniaceae Heliamphoraceae H

elianthaceaeHelianthemaceae Helichrysaceae Heliconiaceae Helicteraceae Heliotropi

aceae Helleboraceae Heloniadaceae HelosidaceaeHelwingiaceae Hemerocallidaceae

Hemimeridaceae Hemiphylacaceae Henriqueziaceae Henslowiaceae Hermanniaceae

Hernandiaceae Herniariaceae Herreriaceae Hesperocallidaceae Heterantheraceae He

teropyxidaceae HeterostylaceaeHibbertiaceae Hibiscaceae Hilleriaceae Himantandra

ceae Hippocastanaceae Hippocrateaceae HippomanaceaeHippophaeaceae Hippurida

ceae Hirtellaceae Holacanthaceae Homaliaceae Hopkinsiaceae Hoplestigmataceae H

ordeaceaeHornschuchiaceae Hortensiaceae Hortoniaceae Hostaceae Houstoniaceae

Huaceae Huerteaceae HugoniaceaeHumbertiaceae Humiriaceae Hyacinthaceae Hyd

atellaceae Hydnoraceae Hydrangeaceae Hydrastidaceae HydrillaceaeHydrocharitacea

e Hydrocotylaceae Hydrogetonaceae Hydroleaceae Hydropeltidaceae Hydrophylacac

eae HydrophyllaceaeHydrostachyaceae Hymenocardiaceae Hypecoaceae Hyperanth

eraceae Hypericaceae Hypopityaceae HypoxidaceaeHypseocharitaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

46

Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta

ceae ImperatoriaceaeInocarpaceae Inulaceae Ionidiaceae Iriarteaceae Iridaceae Irvi

ngiaceae Isnardiaceae Isophysidaceae IteaceaeStrasburgeriaceae Ixiaceae Ixioliriace

ae Ixonanthaceae

Jalapaceae Jamboliferaceae Japonoliriaceae Jasionaceae Jasminiaceae Johnsoniaceae

Joinvilleaceae JuglandaceaeJulianiaceae Juncaceae Juncaginaceae Juniperaceae Jus

siaeaceae Justiciaceae

Kadsuraceae Kaliphoraceae Kalmiaceae Kaniaceae Kiggelariaceae Kingdoniaceae Kin

giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae

Krameriaceae

Labiatae Lacandoniaceae Lachenaliaceae Lacistemataceae Lactoridaceae Lactucaceae

Lagerstroemiaceaee LamiaceaeLampsanaceae Lanariaceae Langsdorffiaceae Lantan

aceae Lapageriaceae Lardizabalaceae Lasiopetalaceae LauraceaeLawsoniaceae Laxm

anniaceae Lecythidaceae Ledaceae Ledocarpaceae Leeaceae Leguminosae Leitneriac

eae LemnaceaeLennoaceae Lentibulariaceae Lentiscaceae Leoniaceae Leonticaceae

Lepidobotryaceae LepidocarpaceaeLepidocaryaceae Lepidocerataceae Leptaulaceae

Leptospermaceae Lepturaceae Lepuropetalaceae LeuctenbergiaceaeLibocedraceae

Licaniaceae Ligustraceae Lilaeaceae Liliaceae Limeaceae Limnanthaceae Limnocharit

aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace

ae Linderniaceae Lindleyaceae Linnaeaceae LippayaceaeLiriaceae Liriodendraceae Li

ssocarpaceae Littorellaceae Loasaceae Lobeliaceaee Loganiaceae LomandraceaeLoni

ceraceae Lopeziaceae Lophiocarpaceae Lophiolaceae Lophiraceae Lophophytaceae L

ophopyxidaceae LoranthaceaeLotaceae Lowiaceae Lupulaceaee Luxemburgiaceae Lu

zuriagaceae Lychnidaceae Lyciaceae LyginiaceaeLygodisodeaceae Lysimachiaceae Ly

thraceae

Macarisiaceae Mackinlayaceae Madiaceae Maesaceae Magnoliaceae Malaceae Male

sherbiaceae MalortieaceaeMalpighiaceae Malvaceae Mangiaceae Manicariaceae Ma

paniaceae Marantaceae Marathraceae MarcgraviaceaeMartyniaceae Mastixiaceae

Matricariaceae Maundiaceae Mayacaceae Medeolaceae Medusagynaceae Peridiscac

eaeMelaleucaceae Melampyraceae Melanophyllaceae Melanthiaceae Melastomatac

eae Meliaceae Melianthaceae MelicaceaeMeliosmaceae Melittidaceae Melochiacea

e Memecylaceae Mendonciaceae Menispermaceae Menthaceae MenyanthaceaeMe

nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

47

Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

Nypaceae Nyssaceae

Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

ceae Papilionaceae Pappophoraceae PapyraceaeParacryphiaceae Parasitaxaceae Par

ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

48

hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

49

cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

50

ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 23: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

23

sequence (see Haston et al 2007) this is of some value Nymphaeales and

Austrobaileyales are here adjacent on the tree but they could be separated by

hundreds of families in the sequence merely by rotating only the node from which

Nymphaeales and their sister group (all other angiosperms minus about 75 species)

arise If adjacent in a book or herbarium or as in these summary pages then it is

relatively easy to relate their characters both apomorphic and plesiomorphic but if

separated by hundreds of pages or two floors in a large building then it is less easy to

get anything from the sequence (Since all orders in the pages of the main part of this

site are preceded by the apomorphies of all nodes immediately below them in the seed

plant phylogeny and because of the linkages that have been built in to the site this

problem is much less serious there there is no sequence) So the particular sequence of

all seed plants that is based on a phylogenetic classification but is to be used in herbaria

or any other place where taxa have to be arranged linearly as in this part of APweb

will be one that maximizes the number of taxa that are both successive branches of the

tree and placed successively in the sequence Since specimens are generally filed under

families the outline of a new family sequence for arranging herbaria and books can now

be suggested (Haston et al 2007) and such a standardized sequence will further help

teaching and learning about plants

ON THINKING ABOUT APOMORPHIES

Identifying apomorphies is important because understanding the evolution of

morphology in the broad sense - ie including anatomy chemistry etc - in the context

of diversification is one of our major goals For identifying apomorphies several

preconditions must be met One needs to have an accurate well supported phylogeny

one has to have examined the right taxa both from the point of view of morphology and

molecules one has to have coded the characters right (ie delimited states

appropriately) and one has to use an appropriate model of evolution when fitting the

variation to the tree (see eg Omland 1999 Stevens 2006b) If the distributions of

apomorphies below are compared with those in other studies (eg Turgeon et al 2001

Bremer et al 2001 Endress 2001 Albach et al 2001a Judd amp Olmstead 2004 D Soltis

et al 2005b Zhang et al 2006 Judd et al 2007) the reader may find a number of

differences Although I have integrated such studies as far as is possible there are five

reasons that link to the preconditions just mentioned why there may be differences

where features are placed on trees

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

24

1 I may not have found all the information about a particular character there may be

disagreement over its interpretation or I have added information

2 The sampling of nearly all molecular studies is very incomplete (see Salisbury amp Kim

2001 for problems caused by sampling) indeed it is commonplace to decry the

incompleteness of molecular sampling But not only is the sampling in molecular studies

often less than we might wish that of the morphological and chemical characters whose

evolution we are interested in understanding is also often very poor So for many

anatomical chemical and embryological characters that are confidently said to

characterise families and other groups we all too often have no idea if those characters

are applicable to the whole clade or just to a subgroup within that clade However as

our sampling improves we can locate changes on the tree more precisely if in

unexpected positions Thus is is sometimes suggested that Ericaceae have ellagic acid

(Soltis et al 2005b) in fact the little evidence we have indicates that only a small clade

of some 80 species has ellagic acid and the rest of the family - itself alone about one

third of all Ericales - does not Similarly Ericaceae are noted for their

ectendomycorrhizal associations but Enkianthus sister to all other Ericaceae

apparently lacks these (Abe 2005) Thus these mycorrhizae and their associated

distinctive hair roots are unlikely to be apomorphies of Ericaceae - however it appears

that other families near Ericaceae may also have ectendomycorrhizae (see Asai 1934)

3 I am fitting characters to a very conservative tree with many polytomies although the

nodes that are utilised are for the most part strongly supported polytomies make the

optimisation of characters that is the assigment of character state change to a

particular node on the tree notably difficult (eg Madison amp Madison 2002) In nearly

all studies of the evolution of characters distributions of characters are optimised on a

more or less fully resolved tree and the construction of supertrees may yield yet more

detailed hypotheses of relationships (for literature on supertrees see Cotton amp

Wilkinson 2007 2008) Of course some nodes on such fully resolved trees andor

supertrees may have little support and optimisations of characters on such trees may

carry correspondingly little conviction Even parts of some of the trees used here have

poor support eg relationships within aquatic Alismatales etc although I indicate

when this is the case

4 Exactly how one goes about optimising a character on a tree is critically important

Even using simple parsimony optimisations (ACCTRAN or DELTRAN ACCelerated

TRANsitions or DELayed TRANsitions) the position of synapomorphies on trees - and

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

25

hence our ideas of evolution - may differ and this is still more true if one uses maximum

likelihood or Bayesian methods and makes apparently reasonable suggestions about

weighting gains over losses (eg Donoghue amp Ackerley 1996 Cunningham et al 1998

Omland 1997 1999 Ree amp Donoghue 1999 Polly 2001 Webster amp Purvis 2001

Ronquist 2004 Crisp amp Cook 2005) Sannier et al (2007) show how in in Arecaceae that

where on a phylogenetic tree one might peg changes in microsporogenesis will depend

on the methods one uses to do this and Pedersen et al (2007) discuss the sometimes

very substantial effect of node support on the posterior probabilities of ancestral

character states Here I use parsimony optimization not always as explicit as it might be

but I have often indicated where there are particularly important uncertainties as to the

positions of particular character changes on the tree

5 Finally although I have paid quite a lot of attention to the delimitation of the

character states that make up all the characterizations I have not spent enough time on

this critical operation If we are interested in understanding evolution then fitting the

basic variation - not character states - to a tree in principle allows greater flexibility in

understanding morphology in the context of local phylogenies (see also Stevens 2000

Endress 2005c) However many character states used here are delimited globally that

is they are circumscribed in the context of the variation shown by individual characters

across all angiosperms andor in the context of classic ideas of character evolution

Character states often have arbitrary limits and serve best to communicate

information whether they are in fact suitable for either phylogenetic analysis or

understanding evolution are separate issues Studies have rather unsurprisingly

perhaps but importantly shown that dividing the one character into different sets of

states may yield differing ideas of evolution of that character (eg Lamb Frye amp Kron

2003 Hibbett 2004) When looking at trees on which character states are optimised

one should bear in mind the problems surrounding the delimitation of states (eg

Stevens 2000 2006b) and the danger of using pollen or other types - constructs

based on many characters that vary independently but which effectively get lost in

these types thus Blackmore et al (2009) decomposed the pollen types in Asteraceae

into 52 characters

I use Remanes three main criteria of homology or better similarity when

determining the basic similarity of structures on different organisms (see Remane 1952)

These criteria are special properties position and intermediates Special properties

include anatomical or chemical characters gene expression data etc Position refers

to the position of an organ with respect to landmarks on the plant Although plants are

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

26

plants and landmarks are not as fixed as one might like trying to work out the relative

positions of parts is a good way of understanding morphology so long as one does not

try to out-do Procrustes Intermediates include intermediates found on the same

plant or on different plants Intermediates may observed during development or be

adult structures Thus one may find changes during development which suggest what

the nature of a particular structure is and very different stuctures on different plants

may turn out to be similar early in development Or one can simply compare different

taxa and see that structures that appear to be very different when comparing two

different taxa can be linked morphologically by looking at other taxa

Nevertheless the use of these criteria may not yield an unambiguous answer as to

what a structure is even given a solid phylogeny and an improved understanding of

development (see Jaramillo amp Kramer 2007 for a useful discussion) As Endress (2005c)

observed a number of features - position function development shape anatomy

histology gene activity and relationships to other taxa that clearly have petals - can be

used to distinguish a petal (for example) from other floral structures if a petal does not

have one of these features is it thereby not a petal Thus Maturen et al (2005) recently

found that floral organ diversity genes (B and C) were expressed in the large white

inflorescence bracts of Cornus (see also Costa et al 2005) Peney et al (2005) noted that

not all monosulcate pollen grains in monocots have the same developmental pathway

and that as a result such pollen might not have the same ancestral state Reeves and

Olmstead (2003) suggested that the genetic mechanisms causing monosymmetry in

Lamiales and Solanales were different and Serna and Martin (2006) described similar

problems with the development of hairs in Arabidopsis when compared with that of

hairs in Antirrhinum and Solanaceae Indeed as one perhaps might expect delimitation

of states and characters does not necessarily become easier with increasing knowledge

of development etc Thus Buzgo et al (2004) Matthews and Endress (2005) and others

have shown how hard it can be to distinguish between eg prophylls and other floral

structures as their behaviour is studied during the course of the development of the

flower Similarly at what concentration is a particular secondary metabolite deemed to

be present (Waterman (2007)

The validity of the approach used here that of fitting morphological variation to a

largely molecular-based tree may be questioned However I think it rather unlikely that

well-supported molecular branches will be overturned by morphological data Indeed

analyses of morphological data alone do provide support for many of the clades evident

in molecule-only analyses and analyses with morphological and molecular data

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

27

together may lead to increased support for clades (eg Hufford 1992 Nandi et al 1998

[but adding morphological data reduces support for a number of critical clades too]

Doyle amp Endress 2000 however in none of these papers is the use of morphology

without ambiguity) It is unfortunately clear that the use of morphology alone may not

suggest problems in the phylogenetic placement of taxa that later turn out to have been

wrongly included (eg Zhang et al 1992) Thus I have been wary of putting much weight

on clades that have only morphological support but note that I have treated molecule-

based clades with low bootstrap or jacknife support values (esp below 70) or low

posterior probabilies (below 095) likewise Although I may have been mistaken in

placing so much emphasis on molecular data in terms of providing the basic

phylogenetic framework for angiosperms morphological and molecular data are only

very rarely in irreconcilably strong conflict There are indeed a few places where the

conflict seems extreme These include the relative positions of the Monimiaceae and

Hernandiaceae (Laurales) the position of Hanguanaceae (Commelinales [as here] or

Zingiberales) and of Triplostegia (is it in Dipsacaceae or Valerianaceae - see

Dipsacales) Fossils are unlikely to affect the topologies of the trees presented here but

see below for their importance in understanding morphological evolution in general and

the evolution of angiosperms in particular Nevertheless some largely reject the idea

that trees based on molecular data alone can recover phylogenetic relationships

especially when branching points are old and prefer to used trees based on analysis of

morphological data including those taken from fossils (Hilton amp Bateman 2006 Farjon

2007)

All in all however the extent of the congruence between morphological and

molecular data is impressive and heartening and many clades can be characterised

morphologically It seemed in 1998 that there were no unambiguous morphological

synapomorphies for angiosperm orders (K Bremer 2000) and this is still true if by

unambiguous is meant non-homoplasious However many orders and other clades

have synapomorphies even if these may be indistinguishable at least at the current

level of morphological and developmental knowledge from parallel occurences

elsewhere As our morphological knowledge increases so too does the number of

apomorphies

Endress and Matthews (2006a) emphasize the importance of tendencies and

developmental constraints when thinking about characters of clades within the rosids -

many characters or character combinations occur in a rather sporadic fashion within

the clade and apparently notably less frequently outside it There are a number of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

28

examples of tendencies here and in the individual order pages such as the features

enclosed by parentheses in groups above the level of family eg see (cuticular waxes

as aggregated rodlets) for the commelinids and the discussion of the distribution of

polyandry that is flowers with numerous stamens in the asterid I + II groups etc

Tendencies may involve more than single characters When thinking of the

associations of features like integument thickness and vascularization of the integument

(Endress amp Matthews 2006a) one can imagine fairly simple developmental

preconditions being operative As Endress and Matthews (2006a) note it is difficult to

imagine vascular tissue developing in an integument only two cells thick Changes here

would be loosely correlated if morphologically linked However the strongly correlated

changes noted by Givnish et al (2005) are ecologically linked but are presumably

morphologicallydevelopmentally independent When there seem to be characters

evolving more or less together tests can be carried out to see if the changes are

concentrated on certain branches of the tree (eg Maddison 1990 Sanderson 1991

Maddison amp Maddison 2000) Such tests have rarely been carried out at higher levels in

angiosperms

One of the most striking examples of a tendency is the distribution of N-fixation

restricted as it is to a monophyletic group of four clades although it has arisen seven (or

perhaps several more) times independently within the N-fixing clade and several

members of two quite different kinds of bacteria are involved (eg Clawson et al 2004

Elliott et al 2007 Sprent amp James 2007) Other examples of work that bears on the issue

of tendencies include the findings that flowers of polysymmetrical Arabidopsis have

genes like TCP1 that are expressed asymmetrically during early development and TCP1

is a probable orthologue of the well-known CYC gene of Antirrhinum that is involved in

the development of monosymmetric slowers there (Cubas et al 2001 Costa et al 2005

etc) Parallelism might build on this underlying morphologically cryptic monosymmetry

even if details of the genetic mechanisms causing the monosymmetry evident in

particular groups may be different (Reeves amp Olmstead 2003 Cubas 2004 see above)

Similarly there have been several recent suggestions that the capability to synthesise a

particular metabolite may be switched off but not lost and so can sometimes be

reacquired (eg Wink amp Witte 1983 Wink 2003 Liscombe et al 2005 Larsson 2007

Waterman 2007) Hence perhaps the rather spotty distribution of many secondary

metabolites like ellagic acid the indole alkaloid camptothecin iridoids etc when

considered in the context of phylogenies As a non-botanical example - but a rather nice

one - Salwini-Plawen and Mayr (1961) suggested some time ago that there has been

considerable parallelism (40-65 or more independent origins) in the evolution of eyes in

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

29

metazoans However Pax 6 seems to be a master control gene that is involved in eye

formation perhaps throughout all bilateralians (eg Gehring amp Ikeo 1999 Erwin amp

Davidson 2002) which changes the nature of the problem Note however that in

general our invocation of evolutionary tendencies to explain the patterns of

distributions of characters is really an admission that we do not understand those

patterns

As we find out more about variation we will probably find fewer and fewer features

constant throughout a group Most unqualified statements of presence and absence

should properly be qualified as usually present or usually absent if one is thinking of

the characterisations on the order pages as encompassing the total variation within a

clade Thus Pistia alone among monocots as so far known has sieve tube plastids with

starch grains not protein crystals However this does not affect the fact that sieve tube

plastids with cuneate protein crystals and lacking starch grains are an apomorphy for

monocots Furthermore fossils in a number of cases suggest character combinations

unknown in extant taxa as may be seen in the discussions of Fagaceae Platanaceae

Iteaceae Calycanthaceae etc Confusing the issue there may be questions as to where

exactly on the tree a particular fossil is to be placed (eg see Nymphaeaceae

Calycanthaceae Archaefructus etc)

To summarize given our current understandings of both phylogenies and

characters evolution of some characters in which we are interested seems very labile

(see eg D Soltis et al 2005b Endress and Matthews 2006a Stevens 2006b Ekman et

al 2008 for a good recent study) and I have been cautious when talking about character

evolution Much effort must continue to be spent in summarizing characters of well-

established clades at all levels providing features by which they may be recognized and

signaling synapomorphies Remember that (1) the basic morphological anatomical and

chemical knowledge of many critical taxa is woefully incomplete (2) different

assumptions about character evolution may greatly affect the position of

synapomorphies on trees (3) in many cases relationships within and between many

groups are too uncertain at present to worry very much about synapomorphies and

(4) we must be clear about what we do and do not not know As mentioned above I

have indicated in a number of places where there are particularly important

uncertainties as to where characters should be placed on the tree Nevertheless it is a

relatively easy matter to update notes such as these and it can be a simple matter to

incorporate new data on characters that have never before been considered in the

context of a tree Much basic - and unfortunately perhaps unfashionable - work must be

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

30

carried to clarify the distribution of morphological anatomical and chemical characters

for examples of what can to be done see the work of P K Endress and collaborators

(floral morphology and development) and S R Jensen and collaborators (iridoids) But

acquiring information about nodal anatomy cell and tissue distribution nodal anatomy

and the like is not difficult at all and should be generally encouraged

But all this is merely a necesary prelude to the understanding of evolution For this

one needs to know a lot more including dating the phylogeny understanding the

function(s) of characters factoring in relevant aspects of the palaeoenvironment in

which an apomorphy first appeared etc

SUMMARY OF THE SYSTEM

Below is a formalised summary of the relationships within orders of the families of seed

plants There are a few families that are not recognised even as options in APG II and

vice versa as well as a few extra orders The families to be recognised in the most

recent edition of Mabberleys The Plant Book (Mabberley 2008) are also largely

consistent with those below But all differences are trivial and will - I hope - eventually

disappear

Square brackets - [] - enclose clades the plus sign - + - designates sister taxa a

comma - - denotes part of a polytomy and quotation marks - - denotes a

paraphyletic group

SEED PLANTS

GYMNOSPERMS

Cycadales

Cycadaceae + Zamiaceae

Ginkgoales

Ginkgoaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

31

Pinales

Pinaceae [[Araucariaceae [Podocarpaceae + Phyllocladaceae]] [Sciadopityaceae

[Taxaceae + Cupressaceae]]]

Gnetales

Ephedraceae [Gnetaceae + Welwitschiaceae]

ANGIOSPERMSFLOWERING PLANTS

Amborellales

Amborellaceae

Nymphaeales

Hydatellaceae [Cabombaceae + Nymphaeaceae]

Austrobaileyales

Austrobaileyaceae [Schisandraceae + Trimeniaceae]

Chloranthales

Chloranthaceae

MAGNOLIIDS

Magnoliales

Myristicaceae [Magnoliaceae [[Himantandraceae + Degeneriaceae] [Eupomatiaceae +

Annonaceae]]]

Laurales

Calycanthaceae [[Siparunaceae [Gomortegaceae + Atherospermataceae]] [Monimiaceae

[Hernandiaceae + Lauraceae]]]

Canellales

Canellaceae + Winteraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

32

Piperales

[Hydnoraceae + Aristolochiaceae] [Piperaceae + Saururaceae]

MONOCOTS

Acorales

Acoraceae

Alismatales

Araceae [Tofieldiaceae [[Alismataceae [Hydrocharitaceae + Butomaceae]]

[Scheuchzeriaceae [Aponogetonaceae [Juncaginaceae [Maundiaceae [[Posidoniaceae

[Ruppiaceae + Cymodoceaceae]] [Zosteraceae + Potamogetonaceae]]]]]]]

Petrosaviales

Petrosaviaceae

Dioscoreales

Nartheciaceae [[Taccaceae + Thismiaceae] [Burmanniaceae + Dioscoreaceae]]

Pandanales

Velloziaceae Triuridaceae Stemonaceae [Pandanaceae + Cyclanthaceae]

Liliales

Corsiaceae [Campynemataceae [Petermanniaceae [Colchicaceae + Alstroemeriaceae]]

Melanthiaceae [[Philesiaceae + Rhipogonaceae] [Smilacaceae + Liliaceae]]]

Asparagales

Orchidaceae [[Boryaceae [Blandfordiaceae [Lanariaceae [Asteliaceae + Hypoxidaceae]]]]

[[Ixioliriaceae + Tecophilaeaceae] [Doryanthaceae [Iridaceae [Xeronemataceae

[Xanthorrhoeaceae [Alliaceae + Asparagaceae]]]]]]]

COMMELINIDS

Unplaced

Dasypogonaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

33

Arecales

Arecaceae

Poales

[Typhaceae + Bromeliaceae] [Rapateaceae [[Thurniaceae [Juncaceae + Cyperaceae]]

[[Mayacaceae [Eriocaulaceae + Xyridaceae]] [Flagellariaceae [Anarthriaceae

[Centrolepidaceae + Restionaceae]]] [Joinvilleaceae [Ecdeiocoleaceae + Poaceae]]]]]]]

Commelinales

[Commelinaceae + Hanguanaceae] [Philydraceae [Haemodoraceae + Pontederiaceae]]

Zingiberales

Musaceae [Strelitziaceae + Lowiaceae] Heliconiaceae [[Cannaceae + Marantaceae]

[Costaceae + Zingiberaceae]]

Ceratophyllales

Ceratophyllaceae

EUDICOTS

Ranunculales

Eupteleaceae [Papaveraceae [[[Lardizabalaceae + Circaeasteraceae] [Menispermaceae

[Berberidaceae + Ranunculaceae]]]]

Sabiales

Sabiaceae

Proteales

Nelumbonaceae [Platanaceae + Proteaceae]

Trochodendrales

Trochodendraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

34

Buxales

Haptanthaceae [Buxaceae + Didymelaceae]

CORE EUDICOTS

Gunnerales

Gunneraceae + Myrothamnaceae

Dilleniales

Dilleniaceae

Saxifragales

Peridiscaceae [[Paeoniaceae [Altingiaceae [Hamamelidaceae [Cercidiphyllaceae +

Daphniphyllaceae]]]] [[Crassulaceae [Aphanopetalaceae [Tetracarpaeaceae

[Penthoraceae + Haloragaceae]]]] [Iteaceae [Grossulariaceae + Saxifragaceae]]]]

Cynomoriaceae unplaced

Vitales

Vitaceae

ROSIDS

FABIDROSID I

Zygophyllales

Krameriaceae + Zygophyllaceae

Celastrales

Lepidobotryaceae + Celastraceae

Oxalidales

Huaceae [[Connaraceae + Oxalidaceae] [Cunoniaceae [Elaeocarpaceae [Brunelliaceae +

Cephalotaceae]]]]

Malpighiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

35

[Achariaceae [Goupiaceae [Violaceae + Passifloraceae] [Lacistemataceae Salicaceae]]

[Lophopyxidaceae Putranjivaceae] [Ctenolophonaceae [Erythroxylaceae +

Rhizophoraceae]] Linaceae Ixonanthaceae Humiriaceae Irvingiaceae

Centroplacaceae Pandaceae [[Bonnetiaceae + Clusiaceae] [Calophyllaceae

[Hypericaceae + Podostemaceae]]] [Malpighiaceae + Elatinaceae] Ochnaceae

[Peraceae [Rafflesiaceae + Euphorbiaceae]] [Phyllanthaceae + Picrodendraceae]

[Balanopaceae [[Trigoniaceae + Dichapetalaceae] [Chrysobalanaceae +

Euphroniaceae]]] Caryocaraceae

N-FIXING CLADE

Fabales

Quillajaceae [Fabaceae [Polygalaceae + Surianaceae]]

Rosales

Rosaceae [[Barbeyaceae [Elaeagnaceae Dirachmaceae Rhamnaceae]] [Ulmaceae

[Cannabaceae [Moraceae + Urticaceae]]]]

Cucurbitales

Anisophylleaceae [[Corynocarpaceae + Coriariaceae] [Cucurbitaceae [Tetramelaceae

[Datiscaceae + Begoniaceae]]]] Apodanthaceae

Fagales

Nothofagaceae [Fagaceae [[Myricaceae + Juglandaceae] [Casuarinaceae

[Ticodendraceae + Betulaceae]]]]

MALVIDROSID II

pgtGeraniales

Geraniaceae [[Melianthaceae + Francoaceae] [Vivianaceae + Ledocarpaceae]]

Myrtales

Combretaceae [[Onagraceae + Lythraceae] [[Vochysiaceae + Myrtaceae]

[Melastomataceae [Crypteroniaceae [Alzateaceae + Penaeaceae]]]]]

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

36

Crossosomatales

[Staphyleaceae [Guamatelaceae [Crossosomataceae + Stachyuraceae]]] [Aphloiaceae

[Geissolomataceae + Strasburgeriaceae]]

Picramniales

Picramniaceae

Sapindales

Biebersteiniaceae Nitrariaceae [Kirkiaceae [Anacardiaceae + Burseraceae]]

[Sapindaceae [Simaroubaceae Rutaceae Meliaceae]]

Huerteales

Gerradinaceae [Dipentodontaceae + Tapisciaceae]

Malvales

Neuradaceae [Thymelaeaceae [Sphaerosepalaceae Bixaceae [Cistaceae

[Sarcolaenaceae + Dipterocarpaceae]] [Cytinaceae + Muntingiaceae] Malvaceae]]

Brassicales

[Akaniaceae + Tropaeolaceae] [[Moringaceae + Caricaceae] [Setchellanthaceae

[Limnanthaceae [[Koeberliniaceae [Bataceae + Salvadoraceae]] [Emblingiaceae

[Pentadiplandraceae [Gyrostemonaceae Resedaceae Stixis and relatives] Tovariaceae

[Capparaceae [Cleomaceae + Brassicaceae]]]]]]]]

Berberidopsidales

Aextoxicaceae + Berberidopsidaceae

Santalales

Erythropalaceae Olacaceae [[Loranthaceae [Misodendraceae + Schoepfiaceae]]

[Opiliaceae + Santalaceae]] Balanophoraceae unplaced

Caryophyllales

[[Droseraceae [Nepenthaceae [Drosophyllaceae [Ancistrocladaceae +

Dioncophylleaceae]]]]] [[Frankeniaceae + Tamaricaceae] [Polygonaceae +

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

37

Plumbaginaceae]]] [Rhabdodendraceae [Simmondsiaceae [[Asteropeiaceae +

Physenaceae] [[Caryophyllaceae [Achatocarpaceae + Amaranthaceae]]

[Stegnospermataceae [Limeaceae [[Lophocarpaceae [Barbeuiaceae [Aizoaceae

[Sarcobataceae Phytolaccaceae Nyctaginaceae]]]] [Molluginaceae [Halophytaceae

Montiaceae Basellaceae Didiereaceae [Talinaceae [Portulacaceae [Anacampseros etc +

Cactaceae]]]]]]]]]]]]]

ASTERIDS

Cornales

[Cornaceae + Nyssaceae] [Hydrangeaceae + Loasaceae] Hydrostachyaceae [Curtisiaceae

+ Grubbiaceae]

Ericales

[Balsaminaceae [Marcgraviaceae + Tetrameristaceae]] [[Polemoniaceae +

Fouquieraceae] Lecythidaceae [[Sladeniaceae + Pentaphylacaceae] [Sapotaceae

[Ebenaceae [Maesaceae [Theophrastaceae [Myrsinaceae + Primulaceae]]]]]

[Mitrastemonaceae Theaceae [Symplocaceae [Styracaceae + Diapensiaceae]]

[[Sarraceniaceae [Roridulaceae + Actinidiaceae] [Clethraceae [Cyrillaceae +

Ericaceae]]]]]]

LAMIIDASTERID I

Unplaced

Metteniusaceae Oncothecaceae Icacinaceae and some unplaced ex-Icacinaceae

Garryales

Garryaceae + Eucommiaceae

Unplaced

Boraginaceae Vahliaceae

Gentianales

Rubiaceae [Gentianaceae [Loganiaceae [Gelsemiaceae + Apocynaceae]]]

Lamiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

38

Plocospermataceae [Oleaceae [Tetrachondraceae [Calceolariaceae

[Gesneriaceae Peltanthera [Pedaliaceae Carlemanniaceae Martyniaceae Stilbaceae

Plantaginaceae Linderniaceae Bignoniaceae Verbenaceae Lamiaceae Paulowniaceae

Schlegeliaceae Thomandersiaceae Phrymaceae Rehmannia Acanthaceae

Scrophulariaceae Orobanchaceae Byblidaceae Lentibulariaceae]]]]]

Solanales

[Montiniaceae [Sphenocleaceae + Hydroleaceae]] [Convolvulaceae + Solanaceae]

CAMPANULIDASTERID II

Aquifoliales

[[Cardiopteridaceae + Stemonuraceae] [Aquifoliaceae [Helwingiaceae +

Phyllonomaceae]]

Asterales

[Rousseaceae + Campanulaceae] Pentaphragmataceae [[Alseuosmiaceae [Phellinaceae

+ Argophyllaceae]] Stylidiaceae [Menyanthaceae [Goodeniaceae [Calyceraceae +

Asteraceae]]]]

Escalloniales

Escalloniaceae

Bruniales

[Bruniaceae + Columelliaceae]

Apiales

Pennantiaceae [Torricelliaceae [Griseliniaceae [Pittosporaceae [Araliaceae

[Myodocarpaceae + Apiaceae]]]]]

Paracryphiales

Paracryphiaceae

Dipsacales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

39

Adoxaceae [Diervillaceae [Caprifoliaceae [Linnaeaceae [Morinaceae [Dipsacaceae +

Valerianaceae]]]]]

LINKS TO ORDERS AND FAMILIES

[Back to Top]

MAIN GROUPINGS

Asterids commelinids core eudicots asterid 1 asterid

2 eudicots gymnosperms Magnoliophyta monocots N-fixing claderosids seed plants

ALPHABETICAL LISTING OF ALL ORDINAL NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | L | M | N | O | P | Q | R | S | T | U | V | W | X | Z

Abietales Acanthales Acerales Acorales Actinidiales Actinostrobales Adoxales Aescul

ales Agavales Aizoales AkanialesAlismatales Alliales Alseuosmiales Alstroemeriales

Altingiales Amaranthales Amaryllidales Amborellales AmbrosialesAmmiales Amomal

es Ancistrocladales Anisophylleales Annonales Anthobolales Apiales Apocynales Apo

nogetonalesAquifoliales Arales Araliales Aralidiales Araucariales Arecales Aristolochi

ales Athrotaxidales Asarales AsclepiadalesAsparagales Asphodelales Asparagales Ast

eliales Atriplicales Aucubales Austrobaileyales Avenales

Balanitales Balanopales Balanophorales Balsaminales Barbeyales Barclayales Batales

Begoniales BerberidalesBerberidopsidales Betulales Biebersteiniales Bignoniales Bix

ales Boraginales Brassicales Brexiales Bromeliales BrunialesBrunoniales Burmannial

es Burserales Butomales Buxales Byblidales

Cactales Callitrichales Calycanthales Calycerales Campanulales Campynematales Can

ellales Cannales CapparalesCaprifoliales Cardiopteridales Carduales Caricales Carle

manniales Caryophyllales Cassiales Casuarinales CelastralesCentrolepidales Cephalot

ales Cephalotaxales Ceratophyllales Cercidiphyllales Chenopodiales Chironiales Chlor

anthalesChrysobalanales Cinchonales Circaeasterales Cistales Citrales Cocosales Col

chicales Columelliales CombretalesCommelinales Connarales Convolvulales Coriarial

es Cornales Corylales Corynocarpales Crassulales CrossosomatalesCucurbitales Cunn

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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inghamiales Cunoniales Cupressales Cycadales Cyclanthales Cymodoceales Cynarales

CynomorialesCyperales Cyrillales Cytinales

Daphnales Daphniphyllales Dasypogonales Datiscales Degeneriales Desfontainiales D

iapensiales Didymelales DillenialesDioales Dioncophyllales Dioscoreales Diospyrales

Dipentodontales Dipsacales Droserales

Ebenales Echiales Elaeagnales Elaeocarpales Elatinales Elodeales Emmotales Empetr

ales Ephedrales Ericales EriocaulalesErythropalaless Escalloniales Eucommiales Euph

orbiales Eupomatiales Eupteleales Euryalaless

Fabales Fagales Falcatifoliales Ficales Flacourtiales Flagellariales Fouquieriales Franc

oales Frangulales

Galiales Garryales Geissolomatales Gentianales Geraniales Gesneriales Ginkgoales G

laucidiales Globulariales GnetalesGoodeniales Greyiales Griseliniales Grossulariales

Grubbiales Gunnerales Gyrocarpales Gyrostemonales

Haemodorales Haloragales Hamamelidales Hanguanales Heisteriales Helleborales He

lwingiales Himantandrales HippuridalesHomaliales Hortensiales Huales Huerteales

Hydatellales Hydnorales Hydrangeales Hydrastidales HydrocharitalesHydropeltidales

Hydrostachyales Hypericales Hypoxidales

Icacinales Illiciales Iridales Irvingales Iteales Ixiales

Jasminales Juglandales Julianiales Juncaginales Juncales

Lacistematales Lactoridales Lamiales Lardizabalales Laurales Lecythidales Ledocarpal

es Leitneriales LentibularialesLigustrales Liliales Limnanthales Linales Liriales Loasal

es Lobeliales Loganiales Lonicerales Loranthales LowialesLythrales

Magnoliales Malpighiales Malvales Marathrales Marcgraviales Mayacales Medusagy

nales Medusandrales MelanthialesMelastomatales Meliales Melianthales Meliosmal

es Menispermales Menyanthales Metteniusales MitrastemonalesMiyoshiales Moni

miales Moringales Musales Myricales Myristicales Myrothamnales Myrsinales Myrta

les

Najadales Nandinales Narcissales Nartheciales Nelumbonales Nepenthales Neuradal

es Nitrariales Nolanales NothofagalesNyctaginales Nymphaeales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Ochnales Oenotherales Olacales Oleales Onagrales Oncothecales Opuntiales Orchid

ales Oxalidales

Saxifragales Pandales Pandanales Papaverales Paracryphiales Parasitaxales Paridales

Parnassiales PassifloralesPenaeales Pennantiales Pentaphragmatales Peridiscales P

etiveriales Petrosaviales Phellinales Philydrales PhyllanthalesPhysenales Phytolaccal

es Picramniales Pinales Pinguiculales Piperales Pittosporales Plantaginales Platanale

sPlumbaginales Poales Podocarpales Podophyllales Podostemales Polemoniales Pol

ygalales Polygonales PontederialesPortulacales Posidoniales Potamogetonales Primu

lales Proteales

Quercales Quillajales Quintiniales

Rafflesiales Ranunculales Rapateales Resedales Restionales Rhabdodendrales Rhamn

ales Rhinanthales RhizophoralesRhodorales Rhoipteleales Roridulales Rosales Rouss

eales Rubiales Ruppiales Rutales

Sabiales Salicales Salvadorales Samolales Samydales Sanguisorbales Santalales Sapin

dales Sapotales SarracenialesSaxegotheales Saxifragales Scheuchzeriales Sciadopitya

les Scleranthales Scrophulariales Scyphostegiales SedalesSilenales Simmondsiales S

milacales Solanales Sphenocleales Sphenostemonales Stangeriales Stellariales Stemo

nalesStilbales Stylidiales Styracales Surianales

Taccales Tamales Tamaricales Taxales Taxodiales Tecophilaeales Terebinthales Tern

stroemiales Theales TheligonalesThymelaeales Tiliales Tofieldiales Torricelliales Tov

ariales Tribelales Trilliales Trimeniales Triuridales TrochodendralesTropaeolales Turn

erales Typhales

Ulmales Urticales

Vacciniales Vahliales Vallisneriales Velloziales Veratrales Verbenales Viburnales Vinc

ales Violales Viscales VitalesVochysiales

Welwitschiales Winterales

Xanthorrhoeales Ximeniales Xyridales

Zamiales Zingiberales Zosterales Zygophyllales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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ALPHABETICAL LISTING OF ALL FAMILY NAMES OF SEED PLANTS WITH LINKS

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In some cases clicking on a family synonym will send you directly to the subfamily in

which the genus of the basionym is to be found

Abietaceae Abolbodaceae Abrophyllaceae Acalyphaceae Acanthaceae [Acanthochla

mydaceae] Acarnaceaee AceraceaeAchariaceae Achatocarpaceae Achradaceae Achy

ranthaceae Acmopylaceae Acoraceae Acristaceae ActaeaceaeActinidiaceae Actinota

ceae Amentotaxaceae Adenogrammaceae Adoxaceae Aegialitidaceae Aegicerataceae

Aegilopaceae Aeginetiaceaee Aegiphilaceae Aesculaceae Aextoxicaceae Agapanthac

eae Agavaceae AgdestidaceaeAgialidaceae Agrimoniaceae Agrostidaceae Ailanthace

ae Aitoniaceae Aizoaceae Akaniaceae Alangiaceae AlchemillaceaeAldrovandaceae A

lismataceae Alliaceae Allioniaceae Allophylaceae Aloaceae Alopecuraceae Alpiniacea

e AlseuosmiaceaeAlsinaceae Alsodeiaceae Alstroemeriaceae Altingiaceae Alzateacea

e Amaranthaceae Amaryllidaceae Amborellaceae Ambrosiaceae Amentotaxaceae A

mmanniaceae Ammiaceae Amomaceae Ampelidopsaceae Amygdalaceae Amyridacea

eAnacardiaceae Anagallidaceae Anarthriaceae Anchusaceae Ancistrocladaceae Andr

omedaceae AndropogonaceaeAndrostachyaceae Androsynaceae Anemarrhenaceae

Anemonaceae Angelicaceae Anisophylleaceae AnnonaceaeAnomochloaceae Anopter

aceae Anrederaceae Anthemidaceae Anthericaceae Anthobolaceae Antidesmataceae

Antirrhinaceae Antoniaceae Aparinaceae Aphanopetalaceae Aphloiaceae Aphyllanth

aceae Apiaceae ApocynaceaeApodanthaceae Aponogetonaceae Aporusaceae Apose

ridaceae Apostasiaceae Aptandraceae AquifoliaceaeAquilariaceae Aquilegiaceae A

raceae Aragoaceae Araliaceae Aralidiaceaee Araucariaceae Arbutaceae Arceuthidace

aeArceuthobiaceae Arctostaphylaceae Arctotidaceae Ardisiaceae Arecaceae Argoph

yllaceae Arisaraceae AristolochiaceaeAristoteliaceae Arjonaceae Armeriaceae Artem

isiaceae Arthrotaxidaceae Artocarpaceae Arundinaceae ArundinellaceaeAsaraceae A

sclepiadaceae Ascyraceaee Asiraceae Asparagaceae Asperulaceae Asphodelaceae As

pidistraceaeeAsteliaceae Asteraceae Asteranthaceae Asteropeiaceae Astragalaceae

Astrocarpaceae AthanasiaceaeAtherospermataceae Atriplicaceae Atropaceae Aucuba

ceae Aurantiaceae Austrobaileyaceae Austrotaxaceae AvenaceaeAverrhoaceae Avet

raceae Avicenniaceae Azaleaceae Azimaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Balanitaceae Balanopaceae Balanophoraceae Balsameaceae Balsaminaceae Bambusa

ceae Barbaceniaceae BarbeuiaceaeBarbeyaceae Barclayaceae Barringtoniaceae Base

llaceae Bataceae Baueraceae Bauhiniaceae Baxteriaceae BegoniaceaeBehniaceae B

elangeraceae Belloniaceae Bembiciaceae Berberidaceae Berberidopsidaceae Berryac

eaee BersamaceaeBertyaceae Berzeliaceae Besleriaceae Betaceae Betulaceae Biebe

rsteiniaceae Bifariaceae Bignoniaceae BischofiaceaeBixaceae Blakeaceae Blandfordia

ceae Blattiaceae Blepharocaryaceae Blitaceae Blyxaceae Boerlagellaceae Bolivaracea

eBombacaceae Bonnetiaceae Bontiaceae Boopidaceae Boraginaceae Borassaceae B

oroniaceae BoryaceaeBotryodendraceae Bougainvilleaceae Boweniaceae Brassicacea

e Brachycaulaceae Bretschneideraceae BrexiaceaeBromeliaceae Brownlowiaceae Br

unelliaceae Bruniaceae Brunoniaceae Brunsvigiaceae Bryoniaceae BuchneraceaeBuci

daceae Buddlejaceae Buglossaceae Bulbocodiaceae Bumeliaceae Bupleuraceae Burc

hardiaceae BurmanniaceaeBurseraceae Butneriaceae Butomaceae Buxaceae Byblida

ceae Byttneriaceae

Cabombaceae Cacaoaceae Cactaceae Caesalpiniaceae Caladiaceae Calamaceae Calce

olariaceae CalectasiaceaeCalendulaceae Callaceae Callicomaceae Calligonaceae Calli

traceae Callitrichaceae Calochortaceae CalophyllaceaeCalthaceae Calycanthaceae Ca

lyceraceae Cambogiaceae Camelliaceae Campanulaceae CampynemataceaeCanacom

yricaceae Canellaceae Cannabaceae Cannaceae Canopodaceae Canotiaceae Cansjera

ceae CapparaceaeCaprariaceae Caprifoliaceae Cardiopteridaceae Carduaceae Carica

ceae Carissaceae Carlemanniaceae CarpinaceaeCarpodetaceae Cartonemataceae Ca

ryocaraceae Caryophyllaceae Casearieaceae Cassiaceae CassipoureaceaeCassythacea

e Castaneaceae Castelaceae Casuarinaceae Catesbaeaceae Cathedraceae Cecropiace

ae CedraceaeCedrelaceae Celastraceae Celosiaceae Celtidaceae Centaureaceae Cen

trolepidaceae Centroplacaceae CepaceaeCephalanthaceae Cephalotaceae Cephalota

xaceae Cerastiaceae Ceratoniaceae Ceratophyllaceae CerberaceaeCercidiphyllaceae

Cercocarpaceae Cercodiaceae Cereaceae Cerinthaceae Ceroxylaceae Cestraceae Cev

alliaceaeChailletiaceae Chamaedoraceae Chamaemoraceae Chamelauciaceae Chaun

ochitonaceae Chelidoniaceae ChelonaceaeChenopodiaceae Chimonanthaceae Chingi

thamnaceae Chionographidaceae Chiranthodendraceae ChironiaceaeChlaenaceae Ch

loanthaceae Chloranthaceae Chloridaceae Chlorogalaceae Chrysobalanaceae Cicerac

eae CichoriaceaeCimicifugaceae Cinchoniaceae Circaeaceae Circaeasteraceae Cissac

eae Cistaceae Citraceae Clematidaceae CleomaceaeClethraceae Clusiaceae Cneorac

eae Cnestidaceae Cnicaceae Cobaeaceae Cochlospermaceae Cocosaceae Coffeaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Colchicaceae Coleogynaceae Columelliaceae Combretaceae Commelinaceae Comocl

adiaceae Compositae CompsoaceaeConantheraceae Connaraceae Conostylidaceae C

onvallariaceae Convolvulaceae Cordiaceae CoreopsidaceaeCoriandraceae Coriariacea

e Coridaceae Corispermaceae Cornaceae Corokiaceae Coronillaceae Correaceae Cor

rigiolaceaeCorsiaceae Corydalaceae Corylaceae Corynocarpaceae Coryphaceae Cost

aceae Cotyledonaceae CoulaceaeCoutariaceae Coutoubiaceae Crassulaceae Crescen

tiaceae Cressaceae Crinaceae Crocaceae CroomiaceaeCrossosomataceae Crotonacea

e Cruciferae Cryptaceae Crypteroniaceae Cryptocorynaceae CryptomeriaceaeCtenol

ophonaceae Cucurbitaceae Cunninghamiaceae Cunoniaceae Cupressaceae Curcumac

eae Curtisiaceae CuscutaceaeCyananthaceae Cyanastraceae Cynanchaceae Cyanella

ceae Cycadaceae Cyclanthaceae CyclantheraceaeCyclocheilaceae Cydoniaceae Cymo

doceaceae Cynaraceae Cynocrambaceae Cynomoriaceae Cyperaceae CyphiaceaeCyp

hocarpaceae Cypripediaceae Cyrillaceae Cyrtandraceae Cyrtanthaceae Cytinaceae

Dacrycarpaceae Dactylanthaceae Damasoniaceae Daphnaceae Daphniphyllaceae Das

ypogonaceae DatiscaceaeDaturaceae Daucaceae Davidiaceae Davidsoniaceae Decais

neaceae Deeringiaceae Degeneriaceae DelphiniaceaeDendrophthoaceae Columelliac

eae Detariaceae Dialypetalanthaceae Dianellaceae Dianthaceae DiapensiaceaeDicha

petalaceae Dichondraceae Diclidantheraceae Dicrastylidiaceae Dictamnaceae Didiere

aceae DidymelaceaeDidymocarpaceae Diegodendraceae Diervillaceae Digitalidaceae

e Dilatridaceae Dilleniaceae Dioaceae DionaeaceaeDioncophyllaceae Dioscoreaceae

Diosmaceae Diospyraceae Dipentodontaceae Diphylleiaceae Diplolaenaceae Dipsaca

ceaeDipterocarpaceae Dirachmaceae Disanthaceae Diselmaceae Dodonaeaceae Do

mbeyaceae Donatiaceae DorsteniaceaeDortmannaceae Doryanthaceae Drabaceae D

racaenaceae Dracontiaceae Drimyidaceae Droseraceae DrosophyllaceaeDryadaceae

Duabangaceae Duckeodendraceae Dulongiaceae Durantaceae Durionaceae Dysphani

aceae

Ebenaceae Eccremidaceae Ecdeiocoleaceae Echiaceae Echinopaceae Ehretiaceae Ela

eagnaceae ElaeocarpaceaeElegiaceae Elatinaceae Ellisiaceae Ellisiophyllaceae Elodea

ceae Elytranthaceae Embeliaceae Emblingiaceae EmmotaceaeEmpetraceae Encepha

lartaceae Engelhardtiaceae Enhalaceae Epacridaceae Ephedraceae Ephemeranthacea

eEpilobiaceae Epimediaceae Eragrostidaceae Eremolepidaceae Eremosynaceae Erica

ceae Erinaceae EriocaulaceaeEriogonaceae Eriospermaceae Erodiaceae Erycibaceae

Eryngiaceae Erysimaceae Erythroniaceae ErythropalaceaeErythrospermaceae Erythro

xylaceae Escalloniaceae Eschscholziaceae Eucomidaceae Eucommiaceae Eucryphiace

aeEuonymaceae Eupatoriaceae Euphorbiaceae Euphrasiaceae Euphroniaceae Eupo

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matiaceae Eupteleaceae EuryalaceaeEustrephaceae Euthemidaceae Eutocaceae Exb

ucklandiaceae Exocarpaceae

Fabaceae Fagaceae Falcatifoliaceae Ferulaceae Festucaceae Ficaceae Ficoidaceae Fit

zroyaceae FlacourtiaceaeFlagellariaceae Flindersiaceae Foetidiaceae Forestieraceae

Fothergillaceae Fouquieriaceae Fragariaceae FrancoaceaeFrangulaceae Frankeniacea

e Fraxinaceae Fraxinellaceae Freycinetiaceae Fritillariaceae Fuchsiaceae Fumariacea

eFunkiaceae

Gaiadendraceae Galacaceae Galanthaceae Galaxiaceae Galedupaceae Galeniaceae G

aliaceae Garciniaceae GardeniaceaeGardneriaceae Garryaceae Geissolomataceae Ge

itonoplesiaceae Gelsemiaceae Geniostomaceae GentianaceaeGeonomataceae Geosir

idaceae Geraniaceae Gerrardinaceae Gesneriaceae Gethyllidaceae Gilliesiaceae Gina

lloaceaeGinkgoaceae Gisekiaceae Gladiolaceae Glaucidiaceae Glechomaceae Glinace

ae Globulariaceae Gnaphaliaceae GnetaceaeGoetzeaceae Gomortegaceae Gomphiac

eae Gomphrenaceae Gonystylaceae Goodeniaceae Gordoniaceae GouaniaceaeGoup

iaceae Gramineae Gratiolaceae Grewiaceae Greyiaceae Grielaceae Griseliniaceae Gr

onoviaceae GrossulariaceaeGrubbiaceae Guaiacanaceae Guamatelaceae Guettardac

eae Gunneraceae Gustaviaceae Guttiferae GyrocarpaceaeGyrostemonaceae

Hachetteaceae Haemanthaceae Haemodoraceae Halesiaceae Hallieraceae Halocarpa

ceae Halophilaceae HalophytaceaeHaloragaceae Hamamelidaceae Hanguanaceae H

aptanthaceae Harmandaceae Hebenstretiaceae HectorellaceaeHederaceae Hedyosm

aceae Hedyotidaceae Hedysaraceae Heisteriaceae Heleniaceae Heliamphoraceae H

elianthaceaeHelianthemaceae Helichrysaceae Heliconiaceae Helicteraceae Heliotropi

aceae Helleboraceae Heloniadaceae HelosidaceaeHelwingiaceae Hemerocallidaceae

Hemimeridaceae Hemiphylacaceae Henriqueziaceae Henslowiaceae Hermanniaceae

Hernandiaceae Herniariaceae Herreriaceae Hesperocallidaceae Heterantheraceae He

teropyxidaceae HeterostylaceaeHibbertiaceae Hibiscaceae Hilleriaceae Himantandra

ceae Hippocastanaceae Hippocrateaceae HippomanaceaeHippophaeaceae Hippurida

ceae Hirtellaceae Holacanthaceae Homaliaceae Hopkinsiaceae Hoplestigmataceae H

ordeaceaeHornschuchiaceae Hortensiaceae Hortoniaceae Hostaceae Houstoniaceae

Huaceae Huerteaceae HugoniaceaeHumbertiaceae Humiriaceae Hyacinthaceae Hyd

atellaceae Hydnoraceae Hydrangeaceae Hydrastidaceae HydrillaceaeHydrocharitacea

e Hydrocotylaceae Hydrogetonaceae Hydroleaceae Hydropeltidaceae Hydrophylacac

eae HydrophyllaceaeHydrostachyaceae Hymenocardiaceae Hypecoaceae Hyperanth

eraceae Hypericaceae Hypopityaceae HypoxidaceaeHypseocharitaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta

ceae ImperatoriaceaeInocarpaceae Inulaceae Ionidiaceae Iriarteaceae Iridaceae Irvi

ngiaceae Isnardiaceae Isophysidaceae IteaceaeStrasburgeriaceae Ixiaceae Ixioliriace

ae Ixonanthaceae

Jalapaceae Jamboliferaceae Japonoliriaceae Jasionaceae Jasminiaceae Johnsoniaceae

Joinvilleaceae JuglandaceaeJulianiaceae Juncaceae Juncaginaceae Juniperaceae Jus

siaeaceae Justiciaceae

Kadsuraceae Kaliphoraceae Kalmiaceae Kaniaceae Kiggelariaceae Kingdoniaceae Kin

giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae

Krameriaceae

Labiatae Lacandoniaceae Lachenaliaceae Lacistemataceae Lactoridaceae Lactucaceae

Lagerstroemiaceaee LamiaceaeLampsanaceae Lanariaceae Langsdorffiaceae Lantan

aceae Lapageriaceae Lardizabalaceae Lasiopetalaceae LauraceaeLawsoniaceae Laxm

anniaceae Lecythidaceae Ledaceae Ledocarpaceae Leeaceae Leguminosae Leitneriac

eae LemnaceaeLennoaceae Lentibulariaceae Lentiscaceae Leoniaceae Leonticaceae

Lepidobotryaceae LepidocarpaceaeLepidocaryaceae Lepidocerataceae Leptaulaceae

Leptospermaceae Lepturaceae Lepuropetalaceae LeuctenbergiaceaeLibocedraceae

Licaniaceae Ligustraceae Lilaeaceae Liliaceae Limeaceae Limnanthaceae Limnocharit

aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace

ae Linderniaceae Lindleyaceae Linnaeaceae LippayaceaeLiriaceae Liriodendraceae Li

ssocarpaceae Littorellaceae Loasaceae Lobeliaceaee Loganiaceae LomandraceaeLoni

ceraceae Lopeziaceae Lophiocarpaceae Lophiolaceae Lophiraceae Lophophytaceae L

ophopyxidaceae LoranthaceaeLotaceae Lowiaceae Lupulaceaee Luxemburgiaceae Lu

zuriagaceae Lychnidaceae Lyciaceae LyginiaceaeLygodisodeaceae Lysimachiaceae Ly

thraceae

Macarisiaceae Mackinlayaceae Madiaceae Maesaceae Magnoliaceae Malaceae Male

sherbiaceae MalortieaceaeMalpighiaceae Malvaceae Mangiaceae Manicariaceae Ma

paniaceae Marantaceae Marathraceae MarcgraviaceaeMartyniaceae Mastixiaceae

Matricariaceae Maundiaceae Mayacaceae Medeolaceae Medusagynaceae Peridiscac

eaeMelaleucaceae Melampyraceae Melanophyllaceae Melanthiaceae Melastomatac

eae Meliaceae Melianthaceae MelicaceaeMeliosmaceae Melittidaceae Melochiacea

e Memecylaceae Mendonciaceae Menispermaceae Menthaceae MenyanthaceaeMe

nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

Nypaceae Nyssaceae

Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

ceae Papilionaceae Pappophoraceae PapyraceaeParacryphiaceae Parasitaxaceae Par

ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

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hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

50

ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 24: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

24

1 I may not have found all the information about a particular character there may be

disagreement over its interpretation or I have added information

2 The sampling of nearly all molecular studies is very incomplete (see Salisbury amp Kim

2001 for problems caused by sampling) indeed it is commonplace to decry the

incompleteness of molecular sampling But not only is the sampling in molecular studies

often less than we might wish that of the morphological and chemical characters whose

evolution we are interested in understanding is also often very poor So for many

anatomical chemical and embryological characters that are confidently said to

characterise families and other groups we all too often have no idea if those characters

are applicable to the whole clade or just to a subgroup within that clade However as

our sampling improves we can locate changes on the tree more precisely if in

unexpected positions Thus is is sometimes suggested that Ericaceae have ellagic acid

(Soltis et al 2005b) in fact the little evidence we have indicates that only a small clade

of some 80 species has ellagic acid and the rest of the family - itself alone about one

third of all Ericales - does not Similarly Ericaceae are noted for their

ectendomycorrhizal associations but Enkianthus sister to all other Ericaceae

apparently lacks these (Abe 2005) Thus these mycorrhizae and their associated

distinctive hair roots are unlikely to be apomorphies of Ericaceae - however it appears

that other families near Ericaceae may also have ectendomycorrhizae (see Asai 1934)

3 I am fitting characters to a very conservative tree with many polytomies although the

nodes that are utilised are for the most part strongly supported polytomies make the

optimisation of characters that is the assigment of character state change to a

particular node on the tree notably difficult (eg Madison amp Madison 2002) In nearly

all studies of the evolution of characters distributions of characters are optimised on a

more or less fully resolved tree and the construction of supertrees may yield yet more

detailed hypotheses of relationships (for literature on supertrees see Cotton amp

Wilkinson 2007 2008) Of course some nodes on such fully resolved trees andor

supertrees may have little support and optimisations of characters on such trees may

carry correspondingly little conviction Even parts of some of the trees used here have

poor support eg relationships within aquatic Alismatales etc although I indicate

when this is the case

4 Exactly how one goes about optimising a character on a tree is critically important

Even using simple parsimony optimisations (ACCTRAN or DELTRAN ACCelerated

TRANsitions or DELayed TRANsitions) the position of synapomorphies on trees - and

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

25

hence our ideas of evolution - may differ and this is still more true if one uses maximum

likelihood or Bayesian methods and makes apparently reasonable suggestions about

weighting gains over losses (eg Donoghue amp Ackerley 1996 Cunningham et al 1998

Omland 1997 1999 Ree amp Donoghue 1999 Polly 2001 Webster amp Purvis 2001

Ronquist 2004 Crisp amp Cook 2005) Sannier et al (2007) show how in in Arecaceae that

where on a phylogenetic tree one might peg changes in microsporogenesis will depend

on the methods one uses to do this and Pedersen et al (2007) discuss the sometimes

very substantial effect of node support on the posterior probabilities of ancestral

character states Here I use parsimony optimization not always as explicit as it might be

but I have often indicated where there are particularly important uncertainties as to the

positions of particular character changes on the tree

5 Finally although I have paid quite a lot of attention to the delimitation of the

character states that make up all the characterizations I have not spent enough time on

this critical operation If we are interested in understanding evolution then fitting the

basic variation - not character states - to a tree in principle allows greater flexibility in

understanding morphology in the context of local phylogenies (see also Stevens 2000

Endress 2005c) However many character states used here are delimited globally that

is they are circumscribed in the context of the variation shown by individual characters

across all angiosperms andor in the context of classic ideas of character evolution

Character states often have arbitrary limits and serve best to communicate

information whether they are in fact suitable for either phylogenetic analysis or

understanding evolution are separate issues Studies have rather unsurprisingly

perhaps but importantly shown that dividing the one character into different sets of

states may yield differing ideas of evolution of that character (eg Lamb Frye amp Kron

2003 Hibbett 2004) When looking at trees on which character states are optimised

one should bear in mind the problems surrounding the delimitation of states (eg

Stevens 2000 2006b) and the danger of using pollen or other types - constructs

based on many characters that vary independently but which effectively get lost in

these types thus Blackmore et al (2009) decomposed the pollen types in Asteraceae

into 52 characters

I use Remanes three main criteria of homology or better similarity when

determining the basic similarity of structures on different organisms (see Remane 1952)

These criteria are special properties position and intermediates Special properties

include anatomical or chemical characters gene expression data etc Position refers

to the position of an organ with respect to landmarks on the plant Although plants are

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

26

plants and landmarks are not as fixed as one might like trying to work out the relative

positions of parts is a good way of understanding morphology so long as one does not

try to out-do Procrustes Intermediates include intermediates found on the same

plant or on different plants Intermediates may observed during development or be

adult structures Thus one may find changes during development which suggest what

the nature of a particular structure is and very different stuctures on different plants

may turn out to be similar early in development Or one can simply compare different

taxa and see that structures that appear to be very different when comparing two

different taxa can be linked morphologically by looking at other taxa

Nevertheless the use of these criteria may not yield an unambiguous answer as to

what a structure is even given a solid phylogeny and an improved understanding of

development (see Jaramillo amp Kramer 2007 for a useful discussion) As Endress (2005c)

observed a number of features - position function development shape anatomy

histology gene activity and relationships to other taxa that clearly have petals - can be

used to distinguish a petal (for example) from other floral structures if a petal does not

have one of these features is it thereby not a petal Thus Maturen et al (2005) recently

found that floral organ diversity genes (B and C) were expressed in the large white

inflorescence bracts of Cornus (see also Costa et al 2005) Peney et al (2005) noted that

not all monosulcate pollen grains in monocots have the same developmental pathway

and that as a result such pollen might not have the same ancestral state Reeves and

Olmstead (2003) suggested that the genetic mechanisms causing monosymmetry in

Lamiales and Solanales were different and Serna and Martin (2006) described similar

problems with the development of hairs in Arabidopsis when compared with that of

hairs in Antirrhinum and Solanaceae Indeed as one perhaps might expect delimitation

of states and characters does not necessarily become easier with increasing knowledge

of development etc Thus Buzgo et al (2004) Matthews and Endress (2005) and others

have shown how hard it can be to distinguish between eg prophylls and other floral

structures as their behaviour is studied during the course of the development of the

flower Similarly at what concentration is a particular secondary metabolite deemed to

be present (Waterman (2007)

The validity of the approach used here that of fitting morphological variation to a

largely molecular-based tree may be questioned However I think it rather unlikely that

well-supported molecular branches will be overturned by morphological data Indeed

analyses of morphological data alone do provide support for many of the clades evident

in molecule-only analyses and analyses with morphological and molecular data

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

27

together may lead to increased support for clades (eg Hufford 1992 Nandi et al 1998

[but adding morphological data reduces support for a number of critical clades too]

Doyle amp Endress 2000 however in none of these papers is the use of morphology

without ambiguity) It is unfortunately clear that the use of morphology alone may not

suggest problems in the phylogenetic placement of taxa that later turn out to have been

wrongly included (eg Zhang et al 1992) Thus I have been wary of putting much weight

on clades that have only morphological support but note that I have treated molecule-

based clades with low bootstrap or jacknife support values (esp below 70) or low

posterior probabilies (below 095) likewise Although I may have been mistaken in

placing so much emphasis on molecular data in terms of providing the basic

phylogenetic framework for angiosperms morphological and molecular data are only

very rarely in irreconcilably strong conflict There are indeed a few places where the

conflict seems extreme These include the relative positions of the Monimiaceae and

Hernandiaceae (Laurales) the position of Hanguanaceae (Commelinales [as here] or

Zingiberales) and of Triplostegia (is it in Dipsacaceae or Valerianaceae - see

Dipsacales) Fossils are unlikely to affect the topologies of the trees presented here but

see below for their importance in understanding morphological evolution in general and

the evolution of angiosperms in particular Nevertheless some largely reject the idea

that trees based on molecular data alone can recover phylogenetic relationships

especially when branching points are old and prefer to used trees based on analysis of

morphological data including those taken from fossils (Hilton amp Bateman 2006 Farjon

2007)

All in all however the extent of the congruence between morphological and

molecular data is impressive and heartening and many clades can be characterised

morphologically It seemed in 1998 that there were no unambiguous morphological

synapomorphies for angiosperm orders (K Bremer 2000) and this is still true if by

unambiguous is meant non-homoplasious However many orders and other clades

have synapomorphies even if these may be indistinguishable at least at the current

level of morphological and developmental knowledge from parallel occurences

elsewhere As our morphological knowledge increases so too does the number of

apomorphies

Endress and Matthews (2006a) emphasize the importance of tendencies and

developmental constraints when thinking about characters of clades within the rosids -

many characters or character combinations occur in a rather sporadic fashion within

the clade and apparently notably less frequently outside it There are a number of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

28

examples of tendencies here and in the individual order pages such as the features

enclosed by parentheses in groups above the level of family eg see (cuticular waxes

as aggregated rodlets) for the commelinids and the discussion of the distribution of

polyandry that is flowers with numerous stamens in the asterid I + II groups etc

Tendencies may involve more than single characters When thinking of the

associations of features like integument thickness and vascularization of the integument

(Endress amp Matthews 2006a) one can imagine fairly simple developmental

preconditions being operative As Endress and Matthews (2006a) note it is difficult to

imagine vascular tissue developing in an integument only two cells thick Changes here

would be loosely correlated if morphologically linked However the strongly correlated

changes noted by Givnish et al (2005) are ecologically linked but are presumably

morphologicallydevelopmentally independent When there seem to be characters

evolving more or less together tests can be carried out to see if the changes are

concentrated on certain branches of the tree (eg Maddison 1990 Sanderson 1991

Maddison amp Maddison 2000) Such tests have rarely been carried out at higher levels in

angiosperms

One of the most striking examples of a tendency is the distribution of N-fixation

restricted as it is to a monophyletic group of four clades although it has arisen seven (or

perhaps several more) times independently within the N-fixing clade and several

members of two quite different kinds of bacteria are involved (eg Clawson et al 2004

Elliott et al 2007 Sprent amp James 2007) Other examples of work that bears on the issue

of tendencies include the findings that flowers of polysymmetrical Arabidopsis have

genes like TCP1 that are expressed asymmetrically during early development and TCP1

is a probable orthologue of the well-known CYC gene of Antirrhinum that is involved in

the development of monosymmetric slowers there (Cubas et al 2001 Costa et al 2005

etc) Parallelism might build on this underlying morphologically cryptic monosymmetry

even if details of the genetic mechanisms causing the monosymmetry evident in

particular groups may be different (Reeves amp Olmstead 2003 Cubas 2004 see above)

Similarly there have been several recent suggestions that the capability to synthesise a

particular metabolite may be switched off but not lost and so can sometimes be

reacquired (eg Wink amp Witte 1983 Wink 2003 Liscombe et al 2005 Larsson 2007

Waterman 2007) Hence perhaps the rather spotty distribution of many secondary

metabolites like ellagic acid the indole alkaloid camptothecin iridoids etc when

considered in the context of phylogenies As a non-botanical example - but a rather nice

one - Salwini-Plawen and Mayr (1961) suggested some time ago that there has been

considerable parallelism (40-65 or more independent origins) in the evolution of eyes in

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

29

metazoans However Pax 6 seems to be a master control gene that is involved in eye

formation perhaps throughout all bilateralians (eg Gehring amp Ikeo 1999 Erwin amp

Davidson 2002) which changes the nature of the problem Note however that in

general our invocation of evolutionary tendencies to explain the patterns of

distributions of characters is really an admission that we do not understand those

patterns

As we find out more about variation we will probably find fewer and fewer features

constant throughout a group Most unqualified statements of presence and absence

should properly be qualified as usually present or usually absent if one is thinking of

the characterisations on the order pages as encompassing the total variation within a

clade Thus Pistia alone among monocots as so far known has sieve tube plastids with

starch grains not protein crystals However this does not affect the fact that sieve tube

plastids with cuneate protein crystals and lacking starch grains are an apomorphy for

monocots Furthermore fossils in a number of cases suggest character combinations

unknown in extant taxa as may be seen in the discussions of Fagaceae Platanaceae

Iteaceae Calycanthaceae etc Confusing the issue there may be questions as to where

exactly on the tree a particular fossil is to be placed (eg see Nymphaeaceae

Calycanthaceae Archaefructus etc)

To summarize given our current understandings of both phylogenies and

characters evolution of some characters in which we are interested seems very labile

(see eg D Soltis et al 2005b Endress and Matthews 2006a Stevens 2006b Ekman et

al 2008 for a good recent study) and I have been cautious when talking about character

evolution Much effort must continue to be spent in summarizing characters of well-

established clades at all levels providing features by which they may be recognized and

signaling synapomorphies Remember that (1) the basic morphological anatomical and

chemical knowledge of many critical taxa is woefully incomplete (2) different

assumptions about character evolution may greatly affect the position of

synapomorphies on trees (3) in many cases relationships within and between many

groups are too uncertain at present to worry very much about synapomorphies and

(4) we must be clear about what we do and do not not know As mentioned above I

have indicated in a number of places where there are particularly important

uncertainties as to where characters should be placed on the tree Nevertheless it is a

relatively easy matter to update notes such as these and it can be a simple matter to

incorporate new data on characters that have never before been considered in the

context of a tree Much basic - and unfortunately perhaps unfashionable - work must be

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

30

carried to clarify the distribution of morphological anatomical and chemical characters

for examples of what can to be done see the work of P K Endress and collaborators

(floral morphology and development) and S R Jensen and collaborators (iridoids) But

acquiring information about nodal anatomy cell and tissue distribution nodal anatomy

and the like is not difficult at all and should be generally encouraged

But all this is merely a necesary prelude to the understanding of evolution For this

one needs to know a lot more including dating the phylogeny understanding the

function(s) of characters factoring in relevant aspects of the palaeoenvironment in

which an apomorphy first appeared etc

SUMMARY OF THE SYSTEM

Below is a formalised summary of the relationships within orders of the families of seed

plants There are a few families that are not recognised even as options in APG II and

vice versa as well as a few extra orders The families to be recognised in the most

recent edition of Mabberleys The Plant Book (Mabberley 2008) are also largely

consistent with those below But all differences are trivial and will - I hope - eventually

disappear

Square brackets - [] - enclose clades the plus sign - + - designates sister taxa a

comma - - denotes part of a polytomy and quotation marks - - denotes a

paraphyletic group

SEED PLANTS

GYMNOSPERMS

Cycadales

Cycadaceae + Zamiaceae

Ginkgoales

Ginkgoaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

31

Pinales

Pinaceae [[Araucariaceae [Podocarpaceae + Phyllocladaceae]] [Sciadopityaceae

[Taxaceae + Cupressaceae]]]

Gnetales

Ephedraceae [Gnetaceae + Welwitschiaceae]

ANGIOSPERMSFLOWERING PLANTS

Amborellales

Amborellaceae

Nymphaeales

Hydatellaceae [Cabombaceae + Nymphaeaceae]

Austrobaileyales

Austrobaileyaceae [Schisandraceae + Trimeniaceae]

Chloranthales

Chloranthaceae

MAGNOLIIDS

Magnoliales

Myristicaceae [Magnoliaceae [[Himantandraceae + Degeneriaceae] [Eupomatiaceae +

Annonaceae]]]

Laurales

Calycanthaceae [[Siparunaceae [Gomortegaceae + Atherospermataceae]] [Monimiaceae

[Hernandiaceae + Lauraceae]]]

Canellales

Canellaceae + Winteraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

32

Piperales

[Hydnoraceae + Aristolochiaceae] [Piperaceae + Saururaceae]

MONOCOTS

Acorales

Acoraceae

Alismatales

Araceae [Tofieldiaceae [[Alismataceae [Hydrocharitaceae + Butomaceae]]

[Scheuchzeriaceae [Aponogetonaceae [Juncaginaceae [Maundiaceae [[Posidoniaceae

[Ruppiaceae + Cymodoceaceae]] [Zosteraceae + Potamogetonaceae]]]]]]]

Petrosaviales

Petrosaviaceae

Dioscoreales

Nartheciaceae [[Taccaceae + Thismiaceae] [Burmanniaceae + Dioscoreaceae]]

Pandanales

Velloziaceae Triuridaceae Stemonaceae [Pandanaceae + Cyclanthaceae]

Liliales

Corsiaceae [Campynemataceae [Petermanniaceae [Colchicaceae + Alstroemeriaceae]]

Melanthiaceae [[Philesiaceae + Rhipogonaceae] [Smilacaceae + Liliaceae]]]

Asparagales

Orchidaceae [[Boryaceae [Blandfordiaceae [Lanariaceae [Asteliaceae + Hypoxidaceae]]]]

[[Ixioliriaceae + Tecophilaeaceae] [Doryanthaceae [Iridaceae [Xeronemataceae

[Xanthorrhoeaceae [Alliaceae + Asparagaceae]]]]]]]

COMMELINIDS

Unplaced

Dasypogonaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Arecales

Arecaceae

Poales

[Typhaceae + Bromeliaceae] [Rapateaceae [[Thurniaceae [Juncaceae + Cyperaceae]]

[[Mayacaceae [Eriocaulaceae + Xyridaceae]] [Flagellariaceae [Anarthriaceae

[Centrolepidaceae + Restionaceae]]] [Joinvilleaceae [Ecdeiocoleaceae + Poaceae]]]]]]]

Commelinales

[Commelinaceae + Hanguanaceae] [Philydraceae [Haemodoraceae + Pontederiaceae]]

Zingiberales

Musaceae [Strelitziaceae + Lowiaceae] Heliconiaceae [[Cannaceae + Marantaceae]

[Costaceae + Zingiberaceae]]

Ceratophyllales

Ceratophyllaceae

EUDICOTS

Ranunculales

Eupteleaceae [Papaveraceae [[[Lardizabalaceae + Circaeasteraceae] [Menispermaceae

[Berberidaceae + Ranunculaceae]]]]

Sabiales

Sabiaceae

Proteales

Nelumbonaceae [Platanaceae + Proteaceae]

Trochodendrales

Trochodendraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

34

Buxales

Haptanthaceae [Buxaceae + Didymelaceae]

CORE EUDICOTS

Gunnerales

Gunneraceae + Myrothamnaceae

Dilleniales

Dilleniaceae

Saxifragales

Peridiscaceae [[Paeoniaceae [Altingiaceae [Hamamelidaceae [Cercidiphyllaceae +

Daphniphyllaceae]]]] [[Crassulaceae [Aphanopetalaceae [Tetracarpaeaceae

[Penthoraceae + Haloragaceae]]]] [Iteaceae [Grossulariaceae + Saxifragaceae]]]]

Cynomoriaceae unplaced

Vitales

Vitaceae

ROSIDS

FABIDROSID I

Zygophyllales

Krameriaceae + Zygophyllaceae

Celastrales

Lepidobotryaceae + Celastraceae

Oxalidales

Huaceae [[Connaraceae + Oxalidaceae] [Cunoniaceae [Elaeocarpaceae [Brunelliaceae +

Cephalotaceae]]]]

Malpighiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

35

[Achariaceae [Goupiaceae [Violaceae + Passifloraceae] [Lacistemataceae Salicaceae]]

[Lophopyxidaceae Putranjivaceae] [Ctenolophonaceae [Erythroxylaceae +

Rhizophoraceae]] Linaceae Ixonanthaceae Humiriaceae Irvingiaceae

Centroplacaceae Pandaceae [[Bonnetiaceae + Clusiaceae] [Calophyllaceae

[Hypericaceae + Podostemaceae]]] [Malpighiaceae + Elatinaceae] Ochnaceae

[Peraceae [Rafflesiaceae + Euphorbiaceae]] [Phyllanthaceae + Picrodendraceae]

[Balanopaceae [[Trigoniaceae + Dichapetalaceae] [Chrysobalanaceae +

Euphroniaceae]]] Caryocaraceae

N-FIXING CLADE

Fabales

Quillajaceae [Fabaceae [Polygalaceae + Surianaceae]]

Rosales

Rosaceae [[Barbeyaceae [Elaeagnaceae Dirachmaceae Rhamnaceae]] [Ulmaceae

[Cannabaceae [Moraceae + Urticaceae]]]]

Cucurbitales

Anisophylleaceae [[Corynocarpaceae + Coriariaceae] [Cucurbitaceae [Tetramelaceae

[Datiscaceae + Begoniaceae]]]] Apodanthaceae

Fagales

Nothofagaceae [Fagaceae [[Myricaceae + Juglandaceae] [Casuarinaceae

[Ticodendraceae + Betulaceae]]]]

MALVIDROSID II

pgtGeraniales

Geraniaceae [[Melianthaceae + Francoaceae] [Vivianaceae + Ledocarpaceae]]

Myrtales

Combretaceae [[Onagraceae + Lythraceae] [[Vochysiaceae + Myrtaceae]

[Melastomataceae [Crypteroniaceae [Alzateaceae + Penaeaceae]]]]]

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

36

Crossosomatales

[Staphyleaceae [Guamatelaceae [Crossosomataceae + Stachyuraceae]]] [Aphloiaceae

[Geissolomataceae + Strasburgeriaceae]]

Picramniales

Picramniaceae

Sapindales

Biebersteiniaceae Nitrariaceae [Kirkiaceae [Anacardiaceae + Burseraceae]]

[Sapindaceae [Simaroubaceae Rutaceae Meliaceae]]

Huerteales

Gerradinaceae [Dipentodontaceae + Tapisciaceae]

Malvales

Neuradaceae [Thymelaeaceae [Sphaerosepalaceae Bixaceae [Cistaceae

[Sarcolaenaceae + Dipterocarpaceae]] [Cytinaceae + Muntingiaceae] Malvaceae]]

Brassicales

[Akaniaceae + Tropaeolaceae] [[Moringaceae + Caricaceae] [Setchellanthaceae

[Limnanthaceae [[Koeberliniaceae [Bataceae + Salvadoraceae]] [Emblingiaceae

[Pentadiplandraceae [Gyrostemonaceae Resedaceae Stixis and relatives] Tovariaceae

[Capparaceae [Cleomaceae + Brassicaceae]]]]]]]]

Berberidopsidales

Aextoxicaceae + Berberidopsidaceae

Santalales

Erythropalaceae Olacaceae [[Loranthaceae [Misodendraceae + Schoepfiaceae]]

[Opiliaceae + Santalaceae]] Balanophoraceae unplaced

Caryophyllales

[[Droseraceae [Nepenthaceae [Drosophyllaceae [Ancistrocladaceae +

Dioncophylleaceae]]]]] [[Frankeniaceae + Tamaricaceae] [Polygonaceae +

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

37

Plumbaginaceae]]] [Rhabdodendraceae [Simmondsiaceae [[Asteropeiaceae +

Physenaceae] [[Caryophyllaceae [Achatocarpaceae + Amaranthaceae]]

[Stegnospermataceae [Limeaceae [[Lophocarpaceae [Barbeuiaceae [Aizoaceae

[Sarcobataceae Phytolaccaceae Nyctaginaceae]]]] [Molluginaceae [Halophytaceae

Montiaceae Basellaceae Didiereaceae [Talinaceae [Portulacaceae [Anacampseros etc +

Cactaceae]]]]]]]]]]]]]

ASTERIDS

Cornales

[Cornaceae + Nyssaceae] [Hydrangeaceae + Loasaceae] Hydrostachyaceae [Curtisiaceae

+ Grubbiaceae]

Ericales

[Balsaminaceae [Marcgraviaceae + Tetrameristaceae]] [[Polemoniaceae +

Fouquieraceae] Lecythidaceae [[Sladeniaceae + Pentaphylacaceae] [Sapotaceae

[Ebenaceae [Maesaceae [Theophrastaceae [Myrsinaceae + Primulaceae]]]]]

[Mitrastemonaceae Theaceae [Symplocaceae [Styracaceae + Diapensiaceae]]

[[Sarraceniaceae [Roridulaceae + Actinidiaceae] [Clethraceae [Cyrillaceae +

Ericaceae]]]]]]

LAMIIDASTERID I

Unplaced

Metteniusaceae Oncothecaceae Icacinaceae and some unplaced ex-Icacinaceae

Garryales

Garryaceae + Eucommiaceae

Unplaced

Boraginaceae Vahliaceae

Gentianales

Rubiaceae [Gentianaceae [Loganiaceae [Gelsemiaceae + Apocynaceae]]]

Lamiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

38

Plocospermataceae [Oleaceae [Tetrachondraceae [Calceolariaceae

[Gesneriaceae Peltanthera [Pedaliaceae Carlemanniaceae Martyniaceae Stilbaceae

Plantaginaceae Linderniaceae Bignoniaceae Verbenaceae Lamiaceae Paulowniaceae

Schlegeliaceae Thomandersiaceae Phrymaceae Rehmannia Acanthaceae

Scrophulariaceae Orobanchaceae Byblidaceae Lentibulariaceae]]]]]

Solanales

[Montiniaceae [Sphenocleaceae + Hydroleaceae]] [Convolvulaceae + Solanaceae]

CAMPANULIDASTERID II

Aquifoliales

[[Cardiopteridaceae + Stemonuraceae] [Aquifoliaceae [Helwingiaceae +

Phyllonomaceae]]

Asterales

[Rousseaceae + Campanulaceae] Pentaphragmataceae [[Alseuosmiaceae [Phellinaceae

+ Argophyllaceae]] Stylidiaceae [Menyanthaceae [Goodeniaceae [Calyceraceae +

Asteraceae]]]]

Escalloniales

Escalloniaceae

Bruniales

[Bruniaceae + Columelliaceae]

Apiales

Pennantiaceae [Torricelliaceae [Griseliniaceae [Pittosporaceae [Araliaceae

[Myodocarpaceae + Apiaceae]]]]]

Paracryphiales

Paracryphiaceae

Dipsacales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

39

Adoxaceae [Diervillaceae [Caprifoliaceae [Linnaeaceae [Morinaceae [Dipsacaceae +

Valerianaceae]]]]]

LINKS TO ORDERS AND FAMILIES

[Back to Top]

MAIN GROUPINGS

Asterids commelinids core eudicots asterid 1 asterid

2 eudicots gymnosperms Magnoliophyta monocots N-fixing claderosids seed plants

ALPHABETICAL LISTING OF ALL ORDINAL NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | L | M | N | O | P | Q | R | S | T | U | V | W | X | Z

Abietales Acanthales Acerales Acorales Actinidiales Actinostrobales Adoxales Aescul

ales Agavales Aizoales AkanialesAlismatales Alliales Alseuosmiales Alstroemeriales

Altingiales Amaranthales Amaryllidales Amborellales AmbrosialesAmmiales Amomal

es Ancistrocladales Anisophylleales Annonales Anthobolales Apiales Apocynales Apo

nogetonalesAquifoliales Arales Araliales Aralidiales Araucariales Arecales Aristolochi

ales Athrotaxidales Asarales AsclepiadalesAsparagales Asphodelales Asparagales Ast

eliales Atriplicales Aucubales Austrobaileyales Avenales

Balanitales Balanopales Balanophorales Balsaminales Barbeyales Barclayales Batales

Begoniales BerberidalesBerberidopsidales Betulales Biebersteiniales Bignoniales Bix

ales Boraginales Brassicales Brexiales Bromeliales BrunialesBrunoniales Burmannial

es Burserales Butomales Buxales Byblidales

Cactales Callitrichales Calycanthales Calycerales Campanulales Campynematales Can

ellales Cannales CapparalesCaprifoliales Cardiopteridales Carduales Caricales Carle

manniales Caryophyllales Cassiales Casuarinales CelastralesCentrolepidales Cephalot

ales Cephalotaxales Ceratophyllales Cercidiphyllales Chenopodiales Chironiales Chlor

anthalesChrysobalanales Cinchonales Circaeasterales Cistales Citrales Cocosales Col

chicales Columelliales CombretalesCommelinales Connarales Convolvulales Coriarial

es Cornales Corylales Corynocarpales Crassulales CrossosomatalesCucurbitales Cunn

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

40

inghamiales Cunoniales Cupressales Cycadales Cyclanthales Cymodoceales Cynarales

CynomorialesCyperales Cyrillales Cytinales

Daphnales Daphniphyllales Dasypogonales Datiscales Degeneriales Desfontainiales D

iapensiales Didymelales DillenialesDioales Dioncophyllales Dioscoreales Diospyrales

Dipentodontales Dipsacales Droserales

Ebenales Echiales Elaeagnales Elaeocarpales Elatinales Elodeales Emmotales Empetr

ales Ephedrales Ericales EriocaulalesErythropalaless Escalloniales Eucommiales Euph

orbiales Eupomatiales Eupteleales Euryalaless

Fabales Fagales Falcatifoliales Ficales Flacourtiales Flagellariales Fouquieriales Franc

oales Frangulales

Galiales Garryales Geissolomatales Gentianales Geraniales Gesneriales Ginkgoales G

laucidiales Globulariales GnetalesGoodeniales Greyiales Griseliniales Grossulariales

Grubbiales Gunnerales Gyrocarpales Gyrostemonales

Haemodorales Haloragales Hamamelidales Hanguanales Heisteriales Helleborales He

lwingiales Himantandrales HippuridalesHomaliales Hortensiales Huales Huerteales

Hydatellales Hydnorales Hydrangeales Hydrastidales HydrocharitalesHydropeltidales

Hydrostachyales Hypericales Hypoxidales

Icacinales Illiciales Iridales Irvingales Iteales Ixiales

Jasminales Juglandales Julianiales Juncaginales Juncales

Lacistematales Lactoridales Lamiales Lardizabalales Laurales Lecythidales Ledocarpal

es Leitneriales LentibularialesLigustrales Liliales Limnanthales Linales Liriales Loasal

es Lobeliales Loganiales Lonicerales Loranthales LowialesLythrales

Magnoliales Malpighiales Malvales Marathrales Marcgraviales Mayacales Medusagy

nales Medusandrales MelanthialesMelastomatales Meliales Melianthales Meliosmal

es Menispermales Menyanthales Metteniusales MitrastemonalesMiyoshiales Moni

miales Moringales Musales Myricales Myristicales Myrothamnales Myrsinales Myrta

les

Najadales Nandinales Narcissales Nartheciales Nelumbonales Nepenthales Neuradal

es Nitrariales Nolanales NothofagalesNyctaginales Nymphaeales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

41

Ochnales Oenotherales Olacales Oleales Onagrales Oncothecales Opuntiales Orchid

ales Oxalidales

Saxifragales Pandales Pandanales Papaverales Paracryphiales Parasitaxales Paridales

Parnassiales PassifloralesPenaeales Pennantiales Pentaphragmatales Peridiscales P

etiveriales Petrosaviales Phellinales Philydrales PhyllanthalesPhysenales Phytolaccal

es Picramniales Pinales Pinguiculales Piperales Pittosporales Plantaginales Platanale

sPlumbaginales Poales Podocarpales Podophyllales Podostemales Polemoniales Pol

ygalales Polygonales PontederialesPortulacales Posidoniales Potamogetonales Primu

lales Proteales

Quercales Quillajales Quintiniales

Rafflesiales Ranunculales Rapateales Resedales Restionales Rhabdodendrales Rhamn

ales Rhinanthales RhizophoralesRhodorales Rhoipteleales Roridulales Rosales Rouss

eales Rubiales Ruppiales Rutales

Sabiales Salicales Salvadorales Samolales Samydales Sanguisorbales Santalales Sapin

dales Sapotales SarracenialesSaxegotheales Saxifragales Scheuchzeriales Sciadopitya

les Scleranthales Scrophulariales Scyphostegiales SedalesSilenales Simmondsiales S

milacales Solanales Sphenocleales Sphenostemonales Stangeriales Stellariales Stemo

nalesStilbales Stylidiales Styracales Surianales

Taccales Tamales Tamaricales Taxales Taxodiales Tecophilaeales Terebinthales Tern

stroemiales Theales TheligonalesThymelaeales Tiliales Tofieldiales Torricelliales Tov

ariales Tribelales Trilliales Trimeniales Triuridales TrochodendralesTropaeolales Turn

erales Typhales

Ulmales Urticales

Vacciniales Vahliales Vallisneriales Velloziales Veratrales Verbenales Viburnales Vinc

ales Violales Viscales VitalesVochysiales

Welwitschiales Winterales

Xanthorrhoeales Ximeniales Xyridales

Zamiales Zingiberales Zosterales Zygophyllales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

42

ALPHABETICAL LISTING OF ALL FAMILY NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | K | L | M | N | O | P | Q | R | S | T | U | V | W | X | Y

|Z

Back to Home

In some cases clicking on a family synonym will send you directly to the subfamily in

which the genus of the basionym is to be found

Abietaceae Abolbodaceae Abrophyllaceae Acalyphaceae Acanthaceae [Acanthochla

mydaceae] Acarnaceaee AceraceaeAchariaceae Achatocarpaceae Achradaceae Achy

ranthaceae Acmopylaceae Acoraceae Acristaceae ActaeaceaeActinidiaceae Actinota

ceae Amentotaxaceae Adenogrammaceae Adoxaceae Aegialitidaceae Aegicerataceae

Aegilopaceae Aeginetiaceaee Aegiphilaceae Aesculaceae Aextoxicaceae Agapanthac

eae Agavaceae AgdestidaceaeAgialidaceae Agrimoniaceae Agrostidaceae Ailanthace

ae Aitoniaceae Aizoaceae Akaniaceae Alangiaceae AlchemillaceaeAldrovandaceae A

lismataceae Alliaceae Allioniaceae Allophylaceae Aloaceae Alopecuraceae Alpiniacea

e AlseuosmiaceaeAlsinaceae Alsodeiaceae Alstroemeriaceae Altingiaceae Alzateacea

e Amaranthaceae Amaryllidaceae Amborellaceae Ambrosiaceae Amentotaxaceae A

mmanniaceae Ammiaceae Amomaceae Ampelidopsaceae Amygdalaceae Amyridacea

eAnacardiaceae Anagallidaceae Anarthriaceae Anchusaceae Ancistrocladaceae Andr

omedaceae AndropogonaceaeAndrostachyaceae Androsynaceae Anemarrhenaceae

Anemonaceae Angelicaceae Anisophylleaceae AnnonaceaeAnomochloaceae Anopter

aceae Anrederaceae Anthemidaceae Anthericaceae Anthobolaceae Antidesmataceae

Antirrhinaceae Antoniaceae Aparinaceae Aphanopetalaceae Aphloiaceae Aphyllanth

aceae Apiaceae ApocynaceaeApodanthaceae Aponogetonaceae Aporusaceae Apose

ridaceae Apostasiaceae Aptandraceae AquifoliaceaeAquilariaceae Aquilegiaceae A

raceae Aragoaceae Araliaceae Aralidiaceaee Araucariaceae Arbutaceae Arceuthidace

aeArceuthobiaceae Arctostaphylaceae Arctotidaceae Ardisiaceae Arecaceae Argoph

yllaceae Arisaraceae AristolochiaceaeAristoteliaceae Arjonaceae Armeriaceae Artem

isiaceae Arthrotaxidaceae Artocarpaceae Arundinaceae ArundinellaceaeAsaraceae A

sclepiadaceae Ascyraceaee Asiraceae Asparagaceae Asperulaceae Asphodelaceae As

pidistraceaeeAsteliaceae Asteraceae Asteranthaceae Asteropeiaceae Astragalaceae

Astrocarpaceae AthanasiaceaeAtherospermataceae Atriplicaceae Atropaceae Aucuba

ceae Aurantiaceae Austrobaileyaceae Austrotaxaceae AvenaceaeAverrhoaceae Avet

raceae Avicenniaceae Azaleaceae Azimaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

43

Balanitaceae Balanopaceae Balanophoraceae Balsameaceae Balsaminaceae Bambusa

ceae Barbaceniaceae BarbeuiaceaeBarbeyaceae Barclayaceae Barringtoniaceae Base

llaceae Bataceae Baueraceae Bauhiniaceae Baxteriaceae BegoniaceaeBehniaceae B

elangeraceae Belloniaceae Bembiciaceae Berberidaceae Berberidopsidaceae Berryac

eaee BersamaceaeBertyaceae Berzeliaceae Besleriaceae Betaceae Betulaceae Biebe

rsteiniaceae Bifariaceae Bignoniaceae BischofiaceaeBixaceae Blakeaceae Blandfordia

ceae Blattiaceae Blepharocaryaceae Blitaceae Blyxaceae Boerlagellaceae Bolivaracea

eBombacaceae Bonnetiaceae Bontiaceae Boopidaceae Boraginaceae Borassaceae B

oroniaceae BoryaceaeBotryodendraceae Bougainvilleaceae Boweniaceae Brassicacea

e Brachycaulaceae Bretschneideraceae BrexiaceaeBromeliaceae Brownlowiaceae Br

unelliaceae Bruniaceae Brunoniaceae Brunsvigiaceae Bryoniaceae BuchneraceaeBuci

daceae Buddlejaceae Buglossaceae Bulbocodiaceae Bumeliaceae Bupleuraceae Burc

hardiaceae BurmanniaceaeBurseraceae Butneriaceae Butomaceae Buxaceae Byblida

ceae Byttneriaceae

Cabombaceae Cacaoaceae Cactaceae Caesalpiniaceae Caladiaceae Calamaceae Calce

olariaceae CalectasiaceaeCalendulaceae Callaceae Callicomaceae Calligonaceae Calli

traceae Callitrichaceae Calochortaceae CalophyllaceaeCalthaceae Calycanthaceae Ca

lyceraceae Cambogiaceae Camelliaceae Campanulaceae CampynemataceaeCanacom

yricaceae Canellaceae Cannabaceae Cannaceae Canopodaceae Canotiaceae Cansjera

ceae CapparaceaeCaprariaceae Caprifoliaceae Cardiopteridaceae Carduaceae Carica

ceae Carissaceae Carlemanniaceae CarpinaceaeCarpodetaceae Cartonemataceae Ca

ryocaraceae Caryophyllaceae Casearieaceae Cassiaceae CassipoureaceaeCassythacea

e Castaneaceae Castelaceae Casuarinaceae Catesbaeaceae Cathedraceae Cecropiace

ae CedraceaeCedrelaceae Celastraceae Celosiaceae Celtidaceae Centaureaceae Cen

trolepidaceae Centroplacaceae CepaceaeCephalanthaceae Cephalotaceae Cephalota

xaceae Cerastiaceae Ceratoniaceae Ceratophyllaceae CerberaceaeCercidiphyllaceae

Cercocarpaceae Cercodiaceae Cereaceae Cerinthaceae Ceroxylaceae Cestraceae Cev

alliaceaeChailletiaceae Chamaedoraceae Chamaemoraceae Chamelauciaceae Chaun

ochitonaceae Chelidoniaceae ChelonaceaeChenopodiaceae Chimonanthaceae Chingi

thamnaceae Chionographidaceae Chiranthodendraceae ChironiaceaeChlaenaceae Ch

loanthaceae Chloranthaceae Chloridaceae Chlorogalaceae Chrysobalanaceae Cicerac

eae CichoriaceaeCimicifugaceae Cinchoniaceae Circaeaceae Circaeasteraceae Cissac

eae Cistaceae Citraceae Clematidaceae CleomaceaeClethraceae Clusiaceae Cneorac

eae Cnestidaceae Cnicaceae Cobaeaceae Cochlospermaceae Cocosaceae Coffeaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

44

Colchicaceae Coleogynaceae Columelliaceae Combretaceae Commelinaceae Comocl

adiaceae Compositae CompsoaceaeConantheraceae Connaraceae Conostylidaceae C

onvallariaceae Convolvulaceae Cordiaceae CoreopsidaceaeCoriandraceae Coriariacea

e Coridaceae Corispermaceae Cornaceae Corokiaceae Coronillaceae Correaceae Cor

rigiolaceaeCorsiaceae Corydalaceae Corylaceae Corynocarpaceae Coryphaceae Cost

aceae Cotyledonaceae CoulaceaeCoutariaceae Coutoubiaceae Crassulaceae Crescen

tiaceae Cressaceae Crinaceae Crocaceae CroomiaceaeCrossosomataceae Crotonacea

e Cruciferae Cryptaceae Crypteroniaceae Cryptocorynaceae CryptomeriaceaeCtenol

ophonaceae Cucurbitaceae Cunninghamiaceae Cunoniaceae Cupressaceae Curcumac

eae Curtisiaceae CuscutaceaeCyananthaceae Cyanastraceae Cynanchaceae Cyanella

ceae Cycadaceae Cyclanthaceae CyclantheraceaeCyclocheilaceae Cydoniaceae Cymo

doceaceae Cynaraceae Cynocrambaceae Cynomoriaceae Cyperaceae CyphiaceaeCyp

hocarpaceae Cypripediaceae Cyrillaceae Cyrtandraceae Cyrtanthaceae Cytinaceae

Dacrycarpaceae Dactylanthaceae Damasoniaceae Daphnaceae Daphniphyllaceae Das

ypogonaceae DatiscaceaeDaturaceae Daucaceae Davidiaceae Davidsoniaceae Decais

neaceae Deeringiaceae Degeneriaceae DelphiniaceaeDendrophthoaceae Columelliac

eae Detariaceae Dialypetalanthaceae Dianellaceae Dianthaceae DiapensiaceaeDicha

petalaceae Dichondraceae Diclidantheraceae Dicrastylidiaceae Dictamnaceae Didiere

aceae DidymelaceaeDidymocarpaceae Diegodendraceae Diervillaceae Digitalidaceae

e Dilatridaceae Dilleniaceae Dioaceae DionaeaceaeDioncophyllaceae Dioscoreaceae

Diosmaceae Diospyraceae Dipentodontaceae Diphylleiaceae Diplolaenaceae Dipsaca

ceaeDipterocarpaceae Dirachmaceae Disanthaceae Diselmaceae Dodonaeaceae Do

mbeyaceae Donatiaceae DorsteniaceaeDortmannaceae Doryanthaceae Drabaceae D

racaenaceae Dracontiaceae Drimyidaceae Droseraceae DrosophyllaceaeDryadaceae

Duabangaceae Duckeodendraceae Dulongiaceae Durantaceae Durionaceae Dysphani

aceae

Ebenaceae Eccremidaceae Ecdeiocoleaceae Echiaceae Echinopaceae Ehretiaceae Ela

eagnaceae ElaeocarpaceaeElegiaceae Elatinaceae Ellisiaceae Ellisiophyllaceae Elodea

ceae Elytranthaceae Embeliaceae Emblingiaceae EmmotaceaeEmpetraceae Encepha

lartaceae Engelhardtiaceae Enhalaceae Epacridaceae Ephedraceae Ephemeranthacea

eEpilobiaceae Epimediaceae Eragrostidaceae Eremolepidaceae Eremosynaceae Erica

ceae Erinaceae EriocaulaceaeEriogonaceae Eriospermaceae Erodiaceae Erycibaceae

Eryngiaceae Erysimaceae Erythroniaceae ErythropalaceaeErythrospermaceae Erythro

xylaceae Escalloniaceae Eschscholziaceae Eucomidaceae Eucommiaceae Eucryphiace

aeEuonymaceae Eupatoriaceae Euphorbiaceae Euphrasiaceae Euphroniaceae Eupo

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

45

matiaceae Eupteleaceae EuryalaceaeEustrephaceae Euthemidaceae Eutocaceae Exb

ucklandiaceae Exocarpaceae

Fabaceae Fagaceae Falcatifoliaceae Ferulaceae Festucaceae Ficaceae Ficoidaceae Fit

zroyaceae FlacourtiaceaeFlagellariaceae Flindersiaceae Foetidiaceae Forestieraceae

Fothergillaceae Fouquieriaceae Fragariaceae FrancoaceaeFrangulaceae Frankeniacea

e Fraxinaceae Fraxinellaceae Freycinetiaceae Fritillariaceae Fuchsiaceae Fumariacea

eFunkiaceae

Gaiadendraceae Galacaceae Galanthaceae Galaxiaceae Galedupaceae Galeniaceae G

aliaceae Garciniaceae GardeniaceaeGardneriaceae Garryaceae Geissolomataceae Ge

itonoplesiaceae Gelsemiaceae Geniostomaceae GentianaceaeGeonomataceae Geosir

idaceae Geraniaceae Gerrardinaceae Gesneriaceae Gethyllidaceae Gilliesiaceae Gina

lloaceaeGinkgoaceae Gisekiaceae Gladiolaceae Glaucidiaceae Glechomaceae Glinace

ae Globulariaceae Gnaphaliaceae GnetaceaeGoetzeaceae Gomortegaceae Gomphiac

eae Gomphrenaceae Gonystylaceae Goodeniaceae Gordoniaceae GouaniaceaeGoup

iaceae Gramineae Gratiolaceae Grewiaceae Greyiaceae Grielaceae Griseliniaceae Gr

onoviaceae GrossulariaceaeGrubbiaceae Guaiacanaceae Guamatelaceae Guettardac

eae Gunneraceae Gustaviaceae Guttiferae GyrocarpaceaeGyrostemonaceae

Hachetteaceae Haemanthaceae Haemodoraceae Halesiaceae Hallieraceae Halocarpa

ceae Halophilaceae HalophytaceaeHaloragaceae Hamamelidaceae Hanguanaceae H

aptanthaceae Harmandaceae Hebenstretiaceae HectorellaceaeHederaceae Hedyosm

aceae Hedyotidaceae Hedysaraceae Heisteriaceae Heleniaceae Heliamphoraceae H

elianthaceaeHelianthemaceae Helichrysaceae Heliconiaceae Helicteraceae Heliotropi

aceae Helleboraceae Heloniadaceae HelosidaceaeHelwingiaceae Hemerocallidaceae

Hemimeridaceae Hemiphylacaceae Henriqueziaceae Henslowiaceae Hermanniaceae

Hernandiaceae Herniariaceae Herreriaceae Hesperocallidaceae Heterantheraceae He

teropyxidaceae HeterostylaceaeHibbertiaceae Hibiscaceae Hilleriaceae Himantandra

ceae Hippocastanaceae Hippocrateaceae HippomanaceaeHippophaeaceae Hippurida

ceae Hirtellaceae Holacanthaceae Homaliaceae Hopkinsiaceae Hoplestigmataceae H

ordeaceaeHornschuchiaceae Hortensiaceae Hortoniaceae Hostaceae Houstoniaceae

Huaceae Huerteaceae HugoniaceaeHumbertiaceae Humiriaceae Hyacinthaceae Hyd

atellaceae Hydnoraceae Hydrangeaceae Hydrastidaceae HydrillaceaeHydrocharitacea

e Hydrocotylaceae Hydrogetonaceae Hydroleaceae Hydropeltidaceae Hydrophylacac

eae HydrophyllaceaeHydrostachyaceae Hymenocardiaceae Hypecoaceae Hyperanth

eraceae Hypericaceae Hypopityaceae HypoxidaceaeHypseocharitaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

46

Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta

ceae ImperatoriaceaeInocarpaceae Inulaceae Ionidiaceae Iriarteaceae Iridaceae Irvi

ngiaceae Isnardiaceae Isophysidaceae IteaceaeStrasburgeriaceae Ixiaceae Ixioliriace

ae Ixonanthaceae

Jalapaceae Jamboliferaceae Japonoliriaceae Jasionaceae Jasminiaceae Johnsoniaceae

Joinvilleaceae JuglandaceaeJulianiaceae Juncaceae Juncaginaceae Juniperaceae Jus

siaeaceae Justiciaceae

Kadsuraceae Kaliphoraceae Kalmiaceae Kaniaceae Kiggelariaceae Kingdoniaceae Kin

giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae

Krameriaceae

Labiatae Lacandoniaceae Lachenaliaceae Lacistemataceae Lactoridaceae Lactucaceae

Lagerstroemiaceaee LamiaceaeLampsanaceae Lanariaceae Langsdorffiaceae Lantan

aceae Lapageriaceae Lardizabalaceae Lasiopetalaceae LauraceaeLawsoniaceae Laxm

anniaceae Lecythidaceae Ledaceae Ledocarpaceae Leeaceae Leguminosae Leitneriac

eae LemnaceaeLennoaceae Lentibulariaceae Lentiscaceae Leoniaceae Leonticaceae

Lepidobotryaceae LepidocarpaceaeLepidocaryaceae Lepidocerataceae Leptaulaceae

Leptospermaceae Lepturaceae Lepuropetalaceae LeuctenbergiaceaeLibocedraceae

Licaniaceae Ligustraceae Lilaeaceae Liliaceae Limeaceae Limnanthaceae Limnocharit

aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace

ae Linderniaceae Lindleyaceae Linnaeaceae LippayaceaeLiriaceae Liriodendraceae Li

ssocarpaceae Littorellaceae Loasaceae Lobeliaceaee Loganiaceae LomandraceaeLoni

ceraceae Lopeziaceae Lophiocarpaceae Lophiolaceae Lophiraceae Lophophytaceae L

ophopyxidaceae LoranthaceaeLotaceae Lowiaceae Lupulaceaee Luxemburgiaceae Lu

zuriagaceae Lychnidaceae Lyciaceae LyginiaceaeLygodisodeaceae Lysimachiaceae Ly

thraceae

Macarisiaceae Mackinlayaceae Madiaceae Maesaceae Magnoliaceae Malaceae Male

sherbiaceae MalortieaceaeMalpighiaceae Malvaceae Mangiaceae Manicariaceae Ma

paniaceae Marantaceae Marathraceae MarcgraviaceaeMartyniaceae Mastixiaceae

Matricariaceae Maundiaceae Mayacaceae Medeolaceae Medusagynaceae Peridiscac

eaeMelaleucaceae Melampyraceae Melanophyllaceae Melanthiaceae Melastomatac

eae Meliaceae Melianthaceae MelicaceaeMeliosmaceae Melittidaceae Melochiacea

e Memecylaceae Mendonciaceae Menispermaceae Menthaceae MenyanthaceaeMe

nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

47

Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

Nypaceae Nyssaceae

Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

ceae Papilionaceae Pappophoraceae PapyraceaeParacryphiaceae Parasitaxaceae Par

ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

48

hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

49

cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

50

ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 25: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

25

hence our ideas of evolution - may differ and this is still more true if one uses maximum

likelihood or Bayesian methods and makes apparently reasonable suggestions about

weighting gains over losses (eg Donoghue amp Ackerley 1996 Cunningham et al 1998

Omland 1997 1999 Ree amp Donoghue 1999 Polly 2001 Webster amp Purvis 2001

Ronquist 2004 Crisp amp Cook 2005) Sannier et al (2007) show how in in Arecaceae that

where on a phylogenetic tree one might peg changes in microsporogenesis will depend

on the methods one uses to do this and Pedersen et al (2007) discuss the sometimes

very substantial effect of node support on the posterior probabilities of ancestral

character states Here I use parsimony optimization not always as explicit as it might be

but I have often indicated where there are particularly important uncertainties as to the

positions of particular character changes on the tree

5 Finally although I have paid quite a lot of attention to the delimitation of the

character states that make up all the characterizations I have not spent enough time on

this critical operation If we are interested in understanding evolution then fitting the

basic variation - not character states - to a tree in principle allows greater flexibility in

understanding morphology in the context of local phylogenies (see also Stevens 2000

Endress 2005c) However many character states used here are delimited globally that

is they are circumscribed in the context of the variation shown by individual characters

across all angiosperms andor in the context of classic ideas of character evolution

Character states often have arbitrary limits and serve best to communicate

information whether they are in fact suitable for either phylogenetic analysis or

understanding evolution are separate issues Studies have rather unsurprisingly

perhaps but importantly shown that dividing the one character into different sets of

states may yield differing ideas of evolution of that character (eg Lamb Frye amp Kron

2003 Hibbett 2004) When looking at trees on which character states are optimised

one should bear in mind the problems surrounding the delimitation of states (eg

Stevens 2000 2006b) and the danger of using pollen or other types - constructs

based on many characters that vary independently but which effectively get lost in

these types thus Blackmore et al (2009) decomposed the pollen types in Asteraceae

into 52 characters

I use Remanes three main criteria of homology or better similarity when

determining the basic similarity of structures on different organisms (see Remane 1952)

These criteria are special properties position and intermediates Special properties

include anatomical or chemical characters gene expression data etc Position refers

to the position of an organ with respect to landmarks on the plant Although plants are

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

26

plants and landmarks are not as fixed as one might like trying to work out the relative

positions of parts is a good way of understanding morphology so long as one does not

try to out-do Procrustes Intermediates include intermediates found on the same

plant or on different plants Intermediates may observed during development or be

adult structures Thus one may find changes during development which suggest what

the nature of a particular structure is and very different stuctures on different plants

may turn out to be similar early in development Or one can simply compare different

taxa and see that structures that appear to be very different when comparing two

different taxa can be linked morphologically by looking at other taxa

Nevertheless the use of these criteria may not yield an unambiguous answer as to

what a structure is even given a solid phylogeny and an improved understanding of

development (see Jaramillo amp Kramer 2007 for a useful discussion) As Endress (2005c)

observed a number of features - position function development shape anatomy

histology gene activity and relationships to other taxa that clearly have petals - can be

used to distinguish a petal (for example) from other floral structures if a petal does not

have one of these features is it thereby not a petal Thus Maturen et al (2005) recently

found that floral organ diversity genes (B and C) were expressed in the large white

inflorescence bracts of Cornus (see also Costa et al 2005) Peney et al (2005) noted that

not all monosulcate pollen grains in monocots have the same developmental pathway

and that as a result such pollen might not have the same ancestral state Reeves and

Olmstead (2003) suggested that the genetic mechanisms causing monosymmetry in

Lamiales and Solanales were different and Serna and Martin (2006) described similar

problems with the development of hairs in Arabidopsis when compared with that of

hairs in Antirrhinum and Solanaceae Indeed as one perhaps might expect delimitation

of states and characters does not necessarily become easier with increasing knowledge

of development etc Thus Buzgo et al (2004) Matthews and Endress (2005) and others

have shown how hard it can be to distinguish between eg prophylls and other floral

structures as their behaviour is studied during the course of the development of the

flower Similarly at what concentration is a particular secondary metabolite deemed to

be present (Waterman (2007)

The validity of the approach used here that of fitting morphological variation to a

largely molecular-based tree may be questioned However I think it rather unlikely that

well-supported molecular branches will be overturned by morphological data Indeed

analyses of morphological data alone do provide support for many of the clades evident

in molecule-only analyses and analyses with morphological and molecular data

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

27

together may lead to increased support for clades (eg Hufford 1992 Nandi et al 1998

[but adding morphological data reduces support for a number of critical clades too]

Doyle amp Endress 2000 however in none of these papers is the use of morphology

without ambiguity) It is unfortunately clear that the use of morphology alone may not

suggest problems in the phylogenetic placement of taxa that later turn out to have been

wrongly included (eg Zhang et al 1992) Thus I have been wary of putting much weight

on clades that have only morphological support but note that I have treated molecule-

based clades with low bootstrap or jacknife support values (esp below 70) or low

posterior probabilies (below 095) likewise Although I may have been mistaken in

placing so much emphasis on molecular data in terms of providing the basic

phylogenetic framework for angiosperms morphological and molecular data are only

very rarely in irreconcilably strong conflict There are indeed a few places where the

conflict seems extreme These include the relative positions of the Monimiaceae and

Hernandiaceae (Laurales) the position of Hanguanaceae (Commelinales [as here] or

Zingiberales) and of Triplostegia (is it in Dipsacaceae or Valerianaceae - see

Dipsacales) Fossils are unlikely to affect the topologies of the trees presented here but

see below for their importance in understanding morphological evolution in general and

the evolution of angiosperms in particular Nevertheless some largely reject the idea

that trees based on molecular data alone can recover phylogenetic relationships

especially when branching points are old and prefer to used trees based on analysis of

morphological data including those taken from fossils (Hilton amp Bateman 2006 Farjon

2007)

All in all however the extent of the congruence between morphological and

molecular data is impressive and heartening and many clades can be characterised

morphologically It seemed in 1998 that there were no unambiguous morphological

synapomorphies for angiosperm orders (K Bremer 2000) and this is still true if by

unambiguous is meant non-homoplasious However many orders and other clades

have synapomorphies even if these may be indistinguishable at least at the current

level of morphological and developmental knowledge from parallel occurences

elsewhere As our morphological knowledge increases so too does the number of

apomorphies

Endress and Matthews (2006a) emphasize the importance of tendencies and

developmental constraints when thinking about characters of clades within the rosids -

many characters or character combinations occur in a rather sporadic fashion within

the clade and apparently notably less frequently outside it There are a number of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

28

examples of tendencies here and in the individual order pages such as the features

enclosed by parentheses in groups above the level of family eg see (cuticular waxes

as aggregated rodlets) for the commelinids and the discussion of the distribution of

polyandry that is flowers with numerous stamens in the asterid I + II groups etc

Tendencies may involve more than single characters When thinking of the

associations of features like integument thickness and vascularization of the integument

(Endress amp Matthews 2006a) one can imagine fairly simple developmental

preconditions being operative As Endress and Matthews (2006a) note it is difficult to

imagine vascular tissue developing in an integument only two cells thick Changes here

would be loosely correlated if morphologically linked However the strongly correlated

changes noted by Givnish et al (2005) are ecologically linked but are presumably

morphologicallydevelopmentally independent When there seem to be characters

evolving more or less together tests can be carried out to see if the changes are

concentrated on certain branches of the tree (eg Maddison 1990 Sanderson 1991

Maddison amp Maddison 2000) Such tests have rarely been carried out at higher levels in

angiosperms

One of the most striking examples of a tendency is the distribution of N-fixation

restricted as it is to a monophyletic group of four clades although it has arisen seven (or

perhaps several more) times independently within the N-fixing clade and several

members of two quite different kinds of bacteria are involved (eg Clawson et al 2004

Elliott et al 2007 Sprent amp James 2007) Other examples of work that bears on the issue

of tendencies include the findings that flowers of polysymmetrical Arabidopsis have

genes like TCP1 that are expressed asymmetrically during early development and TCP1

is a probable orthologue of the well-known CYC gene of Antirrhinum that is involved in

the development of monosymmetric slowers there (Cubas et al 2001 Costa et al 2005

etc) Parallelism might build on this underlying morphologically cryptic monosymmetry

even if details of the genetic mechanisms causing the monosymmetry evident in

particular groups may be different (Reeves amp Olmstead 2003 Cubas 2004 see above)

Similarly there have been several recent suggestions that the capability to synthesise a

particular metabolite may be switched off but not lost and so can sometimes be

reacquired (eg Wink amp Witte 1983 Wink 2003 Liscombe et al 2005 Larsson 2007

Waterman 2007) Hence perhaps the rather spotty distribution of many secondary

metabolites like ellagic acid the indole alkaloid camptothecin iridoids etc when

considered in the context of phylogenies As a non-botanical example - but a rather nice

one - Salwini-Plawen and Mayr (1961) suggested some time ago that there has been

considerable parallelism (40-65 or more independent origins) in the evolution of eyes in

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

29

metazoans However Pax 6 seems to be a master control gene that is involved in eye

formation perhaps throughout all bilateralians (eg Gehring amp Ikeo 1999 Erwin amp

Davidson 2002) which changes the nature of the problem Note however that in

general our invocation of evolutionary tendencies to explain the patterns of

distributions of characters is really an admission that we do not understand those

patterns

As we find out more about variation we will probably find fewer and fewer features

constant throughout a group Most unqualified statements of presence and absence

should properly be qualified as usually present or usually absent if one is thinking of

the characterisations on the order pages as encompassing the total variation within a

clade Thus Pistia alone among monocots as so far known has sieve tube plastids with

starch grains not protein crystals However this does not affect the fact that sieve tube

plastids with cuneate protein crystals and lacking starch grains are an apomorphy for

monocots Furthermore fossils in a number of cases suggest character combinations

unknown in extant taxa as may be seen in the discussions of Fagaceae Platanaceae

Iteaceae Calycanthaceae etc Confusing the issue there may be questions as to where

exactly on the tree a particular fossil is to be placed (eg see Nymphaeaceae

Calycanthaceae Archaefructus etc)

To summarize given our current understandings of both phylogenies and

characters evolution of some characters in which we are interested seems very labile

(see eg D Soltis et al 2005b Endress and Matthews 2006a Stevens 2006b Ekman et

al 2008 for a good recent study) and I have been cautious when talking about character

evolution Much effort must continue to be spent in summarizing characters of well-

established clades at all levels providing features by which they may be recognized and

signaling synapomorphies Remember that (1) the basic morphological anatomical and

chemical knowledge of many critical taxa is woefully incomplete (2) different

assumptions about character evolution may greatly affect the position of

synapomorphies on trees (3) in many cases relationships within and between many

groups are too uncertain at present to worry very much about synapomorphies and

(4) we must be clear about what we do and do not not know As mentioned above I

have indicated in a number of places where there are particularly important

uncertainties as to where characters should be placed on the tree Nevertheless it is a

relatively easy matter to update notes such as these and it can be a simple matter to

incorporate new data on characters that have never before been considered in the

context of a tree Much basic - and unfortunately perhaps unfashionable - work must be

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

30

carried to clarify the distribution of morphological anatomical and chemical characters

for examples of what can to be done see the work of P K Endress and collaborators

(floral morphology and development) and S R Jensen and collaborators (iridoids) But

acquiring information about nodal anatomy cell and tissue distribution nodal anatomy

and the like is not difficult at all and should be generally encouraged

But all this is merely a necesary prelude to the understanding of evolution For this

one needs to know a lot more including dating the phylogeny understanding the

function(s) of characters factoring in relevant aspects of the palaeoenvironment in

which an apomorphy first appeared etc

SUMMARY OF THE SYSTEM

Below is a formalised summary of the relationships within orders of the families of seed

plants There are a few families that are not recognised even as options in APG II and

vice versa as well as a few extra orders The families to be recognised in the most

recent edition of Mabberleys The Plant Book (Mabberley 2008) are also largely

consistent with those below But all differences are trivial and will - I hope - eventually

disappear

Square brackets - [] - enclose clades the plus sign - + - designates sister taxa a

comma - - denotes part of a polytomy and quotation marks - - denotes a

paraphyletic group

SEED PLANTS

GYMNOSPERMS

Cycadales

Cycadaceae + Zamiaceae

Ginkgoales

Ginkgoaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

31

Pinales

Pinaceae [[Araucariaceae [Podocarpaceae + Phyllocladaceae]] [Sciadopityaceae

[Taxaceae + Cupressaceae]]]

Gnetales

Ephedraceae [Gnetaceae + Welwitschiaceae]

ANGIOSPERMSFLOWERING PLANTS

Amborellales

Amborellaceae

Nymphaeales

Hydatellaceae [Cabombaceae + Nymphaeaceae]

Austrobaileyales

Austrobaileyaceae [Schisandraceae + Trimeniaceae]

Chloranthales

Chloranthaceae

MAGNOLIIDS

Magnoliales

Myristicaceae [Magnoliaceae [[Himantandraceae + Degeneriaceae] [Eupomatiaceae +

Annonaceae]]]

Laurales

Calycanthaceae [[Siparunaceae [Gomortegaceae + Atherospermataceae]] [Monimiaceae

[Hernandiaceae + Lauraceae]]]

Canellales

Canellaceae + Winteraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

32

Piperales

[Hydnoraceae + Aristolochiaceae] [Piperaceae + Saururaceae]

MONOCOTS

Acorales

Acoraceae

Alismatales

Araceae [Tofieldiaceae [[Alismataceae [Hydrocharitaceae + Butomaceae]]

[Scheuchzeriaceae [Aponogetonaceae [Juncaginaceae [Maundiaceae [[Posidoniaceae

[Ruppiaceae + Cymodoceaceae]] [Zosteraceae + Potamogetonaceae]]]]]]]

Petrosaviales

Petrosaviaceae

Dioscoreales

Nartheciaceae [[Taccaceae + Thismiaceae] [Burmanniaceae + Dioscoreaceae]]

Pandanales

Velloziaceae Triuridaceae Stemonaceae [Pandanaceae + Cyclanthaceae]

Liliales

Corsiaceae [Campynemataceae [Petermanniaceae [Colchicaceae + Alstroemeriaceae]]

Melanthiaceae [[Philesiaceae + Rhipogonaceae] [Smilacaceae + Liliaceae]]]

Asparagales

Orchidaceae [[Boryaceae [Blandfordiaceae [Lanariaceae [Asteliaceae + Hypoxidaceae]]]]

[[Ixioliriaceae + Tecophilaeaceae] [Doryanthaceae [Iridaceae [Xeronemataceae

[Xanthorrhoeaceae [Alliaceae + Asparagaceae]]]]]]]

COMMELINIDS

Unplaced

Dasypogonaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

33

Arecales

Arecaceae

Poales

[Typhaceae + Bromeliaceae] [Rapateaceae [[Thurniaceae [Juncaceae + Cyperaceae]]

[[Mayacaceae [Eriocaulaceae + Xyridaceae]] [Flagellariaceae [Anarthriaceae

[Centrolepidaceae + Restionaceae]]] [Joinvilleaceae [Ecdeiocoleaceae + Poaceae]]]]]]]

Commelinales

[Commelinaceae + Hanguanaceae] [Philydraceae [Haemodoraceae + Pontederiaceae]]

Zingiberales

Musaceae [Strelitziaceae + Lowiaceae] Heliconiaceae [[Cannaceae + Marantaceae]

[Costaceae + Zingiberaceae]]

Ceratophyllales

Ceratophyllaceae

EUDICOTS

Ranunculales

Eupteleaceae [Papaveraceae [[[Lardizabalaceae + Circaeasteraceae] [Menispermaceae

[Berberidaceae + Ranunculaceae]]]]

Sabiales

Sabiaceae

Proteales

Nelumbonaceae [Platanaceae + Proteaceae]

Trochodendrales

Trochodendraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

34

Buxales

Haptanthaceae [Buxaceae + Didymelaceae]

CORE EUDICOTS

Gunnerales

Gunneraceae + Myrothamnaceae

Dilleniales

Dilleniaceae

Saxifragales

Peridiscaceae [[Paeoniaceae [Altingiaceae [Hamamelidaceae [Cercidiphyllaceae +

Daphniphyllaceae]]]] [[Crassulaceae [Aphanopetalaceae [Tetracarpaeaceae

[Penthoraceae + Haloragaceae]]]] [Iteaceae [Grossulariaceae + Saxifragaceae]]]]

Cynomoriaceae unplaced

Vitales

Vitaceae

ROSIDS

FABIDROSID I

Zygophyllales

Krameriaceae + Zygophyllaceae

Celastrales

Lepidobotryaceae + Celastraceae

Oxalidales

Huaceae [[Connaraceae + Oxalidaceae] [Cunoniaceae [Elaeocarpaceae [Brunelliaceae +

Cephalotaceae]]]]

Malpighiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

35

[Achariaceae [Goupiaceae [Violaceae + Passifloraceae] [Lacistemataceae Salicaceae]]

[Lophopyxidaceae Putranjivaceae] [Ctenolophonaceae [Erythroxylaceae +

Rhizophoraceae]] Linaceae Ixonanthaceae Humiriaceae Irvingiaceae

Centroplacaceae Pandaceae [[Bonnetiaceae + Clusiaceae] [Calophyllaceae

[Hypericaceae + Podostemaceae]]] [Malpighiaceae + Elatinaceae] Ochnaceae

[Peraceae [Rafflesiaceae + Euphorbiaceae]] [Phyllanthaceae + Picrodendraceae]

[Balanopaceae [[Trigoniaceae + Dichapetalaceae] [Chrysobalanaceae +

Euphroniaceae]]] Caryocaraceae

N-FIXING CLADE

Fabales

Quillajaceae [Fabaceae [Polygalaceae + Surianaceae]]

Rosales

Rosaceae [[Barbeyaceae [Elaeagnaceae Dirachmaceae Rhamnaceae]] [Ulmaceae

[Cannabaceae [Moraceae + Urticaceae]]]]

Cucurbitales

Anisophylleaceae [[Corynocarpaceae + Coriariaceae] [Cucurbitaceae [Tetramelaceae

[Datiscaceae + Begoniaceae]]]] Apodanthaceae

Fagales

Nothofagaceae [Fagaceae [[Myricaceae + Juglandaceae] [Casuarinaceae

[Ticodendraceae + Betulaceae]]]]

MALVIDROSID II

pgtGeraniales

Geraniaceae [[Melianthaceae + Francoaceae] [Vivianaceae + Ledocarpaceae]]

Myrtales

Combretaceae [[Onagraceae + Lythraceae] [[Vochysiaceae + Myrtaceae]

[Melastomataceae [Crypteroniaceae [Alzateaceae + Penaeaceae]]]]]

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

36

Crossosomatales

[Staphyleaceae [Guamatelaceae [Crossosomataceae + Stachyuraceae]]] [Aphloiaceae

[Geissolomataceae + Strasburgeriaceae]]

Picramniales

Picramniaceae

Sapindales

Biebersteiniaceae Nitrariaceae [Kirkiaceae [Anacardiaceae + Burseraceae]]

[Sapindaceae [Simaroubaceae Rutaceae Meliaceae]]

Huerteales

Gerradinaceae [Dipentodontaceae + Tapisciaceae]

Malvales

Neuradaceae [Thymelaeaceae [Sphaerosepalaceae Bixaceae [Cistaceae

[Sarcolaenaceae + Dipterocarpaceae]] [Cytinaceae + Muntingiaceae] Malvaceae]]

Brassicales

[Akaniaceae + Tropaeolaceae] [[Moringaceae + Caricaceae] [Setchellanthaceae

[Limnanthaceae [[Koeberliniaceae [Bataceae + Salvadoraceae]] [Emblingiaceae

[Pentadiplandraceae [Gyrostemonaceae Resedaceae Stixis and relatives] Tovariaceae

[Capparaceae [Cleomaceae + Brassicaceae]]]]]]]]

Berberidopsidales

Aextoxicaceae + Berberidopsidaceae

Santalales

Erythropalaceae Olacaceae [[Loranthaceae [Misodendraceae + Schoepfiaceae]]

[Opiliaceae + Santalaceae]] Balanophoraceae unplaced

Caryophyllales

[[Droseraceae [Nepenthaceae [Drosophyllaceae [Ancistrocladaceae +

Dioncophylleaceae]]]]] [[Frankeniaceae + Tamaricaceae] [Polygonaceae +

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

37

Plumbaginaceae]]] [Rhabdodendraceae [Simmondsiaceae [[Asteropeiaceae +

Physenaceae] [[Caryophyllaceae [Achatocarpaceae + Amaranthaceae]]

[Stegnospermataceae [Limeaceae [[Lophocarpaceae [Barbeuiaceae [Aizoaceae

[Sarcobataceae Phytolaccaceae Nyctaginaceae]]]] [Molluginaceae [Halophytaceae

Montiaceae Basellaceae Didiereaceae [Talinaceae [Portulacaceae [Anacampseros etc +

Cactaceae]]]]]]]]]]]]]

ASTERIDS

Cornales

[Cornaceae + Nyssaceae] [Hydrangeaceae + Loasaceae] Hydrostachyaceae [Curtisiaceae

+ Grubbiaceae]

Ericales

[Balsaminaceae [Marcgraviaceae + Tetrameristaceae]] [[Polemoniaceae +

Fouquieraceae] Lecythidaceae [[Sladeniaceae + Pentaphylacaceae] [Sapotaceae

[Ebenaceae [Maesaceae [Theophrastaceae [Myrsinaceae + Primulaceae]]]]]

[Mitrastemonaceae Theaceae [Symplocaceae [Styracaceae + Diapensiaceae]]

[[Sarraceniaceae [Roridulaceae + Actinidiaceae] [Clethraceae [Cyrillaceae +

Ericaceae]]]]]]

LAMIIDASTERID I

Unplaced

Metteniusaceae Oncothecaceae Icacinaceae and some unplaced ex-Icacinaceae

Garryales

Garryaceae + Eucommiaceae

Unplaced

Boraginaceae Vahliaceae

Gentianales

Rubiaceae [Gentianaceae [Loganiaceae [Gelsemiaceae + Apocynaceae]]]

Lamiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

38

Plocospermataceae [Oleaceae [Tetrachondraceae [Calceolariaceae

[Gesneriaceae Peltanthera [Pedaliaceae Carlemanniaceae Martyniaceae Stilbaceae

Plantaginaceae Linderniaceae Bignoniaceae Verbenaceae Lamiaceae Paulowniaceae

Schlegeliaceae Thomandersiaceae Phrymaceae Rehmannia Acanthaceae

Scrophulariaceae Orobanchaceae Byblidaceae Lentibulariaceae]]]]]

Solanales

[Montiniaceae [Sphenocleaceae + Hydroleaceae]] [Convolvulaceae + Solanaceae]

CAMPANULIDASTERID II

Aquifoliales

[[Cardiopteridaceae + Stemonuraceae] [Aquifoliaceae [Helwingiaceae +

Phyllonomaceae]]

Asterales

[Rousseaceae + Campanulaceae] Pentaphragmataceae [[Alseuosmiaceae [Phellinaceae

+ Argophyllaceae]] Stylidiaceae [Menyanthaceae [Goodeniaceae [Calyceraceae +

Asteraceae]]]]

Escalloniales

Escalloniaceae

Bruniales

[Bruniaceae + Columelliaceae]

Apiales

Pennantiaceae [Torricelliaceae [Griseliniaceae [Pittosporaceae [Araliaceae

[Myodocarpaceae + Apiaceae]]]]]

Paracryphiales

Paracryphiaceae

Dipsacales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

39

Adoxaceae [Diervillaceae [Caprifoliaceae [Linnaeaceae [Morinaceae [Dipsacaceae +

Valerianaceae]]]]]

LINKS TO ORDERS AND FAMILIES

[Back to Top]

MAIN GROUPINGS

Asterids commelinids core eudicots asterid 1 asterid

2 eudicots gymnosperms Magnoliophyta monocots N-fixing claderosids seed plants

ALPHABETICAL LISTING OF ALL ORDINAL NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | L | M | N | O | P | Q | R | S | T | U | V | W | X | Z

Abietales Acanthales Acerales Acorales Actinidiales Actinostrobales Adoxales Aescul

ales Agavales Aizoales AkanialesAlismatales Alliales Alseuosmiales Alstroemeriales

Altingiales Amaranthales Amaryllidales Amborellales AmbrosialesAmmiales Amomal

es Ancistrocladales Anisophylleales Annonales Anthobolales Apiales Apocynales Apo

nogetonalesAquifoliales Arales Araliales Aralidiales Araucariales Arecales Aristolochi

ales Athrotaxidales Asarales AsclepiadalesAsparagales Asphodelales Asparagales Ast

eliales Atriplicales Aucubales Austrobaileyales Avenales

Balanitales Balanopales Balanophorales Balsaminales Barbeyales Barclayales Batales

Begoniales BerberidalesBerberidopsidales Betulales Biebersteiniales Bignoniales Bix

ales Boraginales Brassicales Brexiales Bromeliales BrunialesBrunoniales Burmannial

es Burserales Butomales Buxales Byblidales

Cactales Callitrichales Calycanthales Calycerales Campanulales Campynematales Can

ellales Cannales CapparalesCaprifoliales Cardiopteridales Carduales Caricales Carle

manniales Caryophyllales Cassiales Casuarinales CelastralesCentrolepidales Cephalot

ales Cephalotaxales Ceratophyllales Cercidiphyllales Chenopodiales Chironiales Chlor

anthalesChrysobalanales Cinchonales Circaeasterales Cistales Citrales Cocosales Col

chicales Columelliales CombretalesCommelinales Connarales Convolvulales Coriarial

es Cornales Corylales Corynocarpales Crassulales CrossosomatalesCucurbitales Cunn

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

40

inghamiales Cunoniales Cupressales Cycadales Cyclanthales Cymodoceales Cynarales

CynomorialesCyperales Cyrillales Cytinales

Daphnales Daphniphyllales Dasypogonales Datiscales Degeneriales Desfontainiales D

iapensiales Didymelales DillenialesDioales Dioncophyllales Dioscoreales Diospyrales

Dipentodontales Dipsacales Droserales

Ebenales Echiales Elaeagnales Elaeocarpales Elatinales Elodeales Emmotales Empetr

ales Ephedrales Ericales EriocaulalesErythropalaless Escalloniales Eucommiales Euph

orbiales Eupomatiales Eupteleales Euryalaless

Fabales Fagales Falcatifoliales Ficales Flacourtiales Flagellariales Fouquieriales Franc

oales Frangulales

Galiales Garryales Geissolomatales Gentianales Geraniales Gesneriales Ginkgoales G

laucidiales Globulariales GnetalesGoodeniales Greyiales Griseliniales Grossulariales

Grubbiales Gunnerales Gyrocarpales Gyrostemonales

Haemodorales Haloragales Hamamelidales Hanguanales Heisteriales Helleborales He

lwingiales Himantandrales HippuridalesHomaliales Hortensiales Huales Huerteales

Hydatellales Hydnorales Hydrangeales Hydrastidales HydrocharitalesHydropeltidales

Hydrostachyales Hypericales Hypoxidales

Icacinales Illiciales Iridales Irvingales Iteales Ixiales

Jasminales Juglandales Julianiales Juncaginales Juncales

Lacistematales Lactoridales Lamiales Lardizabalales Laurales Lecythidales Ledocarpal

es Leitneriales LentibularialesLigustrales Liliales Limnanthales Linales Liriales Loasal

es Lobeliales Loganiales Lonicerales Loranthales LowialesLythrales

Magnoliales Malpighiales Malvales Marathrales Marcgraviales Mayacales Medusagy

nales Medusandrales MelanthialesMelastomatales Meliales Melianthales Meliosmal

es Menispermales Menyanthales Metteniusales MitrastemonalesMiyoshiales Moni

miales Moringales Musales Myricales Myristicales Myrothamnales Myrsinales Myrta

les

Najadales Nandinales Narcissales Nartheciales Nelumbonales Nepenthales Neuradal

es Nitrariales Nolanales NothofagalesNyctaginales Nymphaeales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

41

Ochnales Oenotherales Olacales Oleales Onagrales Oncothecales Opuntiales Orchid

ales Oxalidales

Saxifragales Pandales Pandanales Papaverales Paracryphiales Parasitaxales Paridales

Parnassiales PassifloralesPenaeales Pennantiales Pentaphragmatales Peridiscales P

etiveriales Petrosaviales Phellinales Philydrales PhyllanthalesPhysenales Phytolaccal

es Picramniales Pinales Pinguiculales Piperales Pittosporales Plantaginales Platanale

sPlumbaginales Poales Podocarpales Podophyllales Podostemales Polemoniales Pol

ygalales Polygonales PontederialesPortulacales Posidoniales Potamogetonales Primu

lales Proteales

Quercales Quillajales Quintiniales

Rafflesiales Ranunculales Rapateales Resedales Restionales Rhabdodendrales Rhamn

ales Rhinanthales RhizophoralesRhodorales Rhoipteleales Roridulales Rosales Rouss

eales Rubiales Ruppiales Rutales

Sabiales Salicales Salvadorales Samolales Samydales Sanguisorbales Santalales Sapin

dales Sapotales SarracenialesSaxegotheales Saxifragales Scheuchzeriales Sciadopitya

les Scleranthales Scrophulariales Scyphostegiales SedalesSilenales Simmondsiales S

milacales Solanales Sphenocleales Sphenostemonales Stangeriales Stellariales Stemo

nalesStilbales Stylidiales Styracales Surianales

Taccales Tamales Tamaricales Taxales Taxodiales Tecophilaeales Terebinthales Tern

stroemiales Theales TheligonalesThymelaeales Tiliales Tofieldiales Torricelliales Tov

ariales Tribelales Trilliales Trimeniales Triuridales TrochodendralesTropaeolales Turn

erales Typhales

Ulmales Urticales

Vacciniales Vahliales Vallisneriales Velloziales Veratrales Verbenales Viburnales Vinc

ales Violales Viscales VitalesVochysiales

Welwitschiales Winterales

Xanthorrhoeales Ximeniales Xyridales

Zamiales Zingiberales Zosterales Zygophyllales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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ALPHABETICAL LISTING OF ALL FAMILY NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | K | L | M | N | O | P | Q | R | S | T | U | V | W | X | Y

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In some cases clicking on a family synonym will send you directly to the subfamily in

which the genus of the basionym is to be found

Abietaceae Abolbodaceae Abrophyllaceae Acalyphaceae Acanthaceae [Acanthochla

mydaceae] Acarnaceaee AceraceaeAchariaceae Achatocarpaceae Achradaceae Achy

ranthaceae Acmopylaceae Acoraceae Acristaceae ActaeaceaeActinidiaceae Actinota

ceae Amentotaxaceae Adenogrammaceae Adoxaceae Aegialitidaceae Aegicerataceae

Aegilopaceae Aeginetiaceaee Aegiphilaceae Aesculaceae Aextoxicaceae Agapanthac

eae Agavaceae AgdestidaceaeAgialidaceae Agrimoniaceae Agrostidaceae Ailanthace

ae Aitoniaceae Aizoaceae Akaniaceae Alangiaceae AlchemillaceaeAldrovandaceae A

lismataceae Alliaceae Allioniaceae Allophylaceae Aloaceae Alopecuraceae Alpiniacea

e AlseuosmiaceaeAlsinaceae Alsodeiaceae Alstroemeriaceae Altingiaceae Alzateacea

e Amaranthaceae Amaryllidaceae Amborellaceae Ambrosiaceae Amentotaxaceae A

mmanniaceae Ammiaceae Amomaceae Ampelidopsaceae Amygdalaceae Amyridacea

eAnacardiaceae Anagallidaceae Anarthriaceae Anchusaceae Ancistrocladaceae Andr

omedaceae AndropogonaceaeAndrostachyaceae Androsynaceae Anemarrhenaceae

Anemonaceae Angelicaceae Anisophylleaceae AnnonaceaeAnomochloaceae Anopter

aceae Anrederaceae Anthemidaceae Anthericaceae Anthobolaceae Antidesmataceae

Antirrhinaceae Antoniaceae Aparinaceae Aphanopetalaceae Aphloiaceae Aphyllanth

aceae Apiaceae ApocynaceaeApodanthaceae Aponogetonaceae Aporusaceae Apose

ridaceae Apostasiaceae Aptandraceae AquifoliaceaeAquilariaceae Aquilegiaceae A

raceae Aragoaceae Araliaceae Aralidiaceaee Araucariaceae Arbutaceae Arceuthidace

aeArceuthobiaceae Arctostaphylaceae Arctotidaceae Ardisiaceae Arecaceae Argoph

yllaceae Arisaraceae AristolochiaceaeAristoteliaceae Arjonaceae Armeriaceae Artem

isiaceae Arthrotaxidaceae Artocarpaceae Arundinaceae ArundinellaceaeAsaraceae A

sclepiadaceae Ascyraceaee Asiraceae Asparagaceae Asperulaceae Asphodelaceae As

pidistraceaeeAsteliaceae Asteraceae Asteranthaceae Asteropeiaceae Astragalaceae

Astrocarpaceae AthanasiaceaeAtherospermataceae Atriplicaceae Atropaceae Aucuba

ceae Aurantiaceae Austrobaileyaceae Austrotaxaceae AvenaceaeAverrhoaceae Avet

raceae Avicenniaceae Azaleaceae Azimaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Balanitaceae Balanopaceae Balanophoraceae Balsameaceae Balsaminaceae Bambusa

ceae Barbaceniaceae BarbeuiaceaeBarbeyaceae Barclayaceae Barringtoniaceae Base

llaceae Bataceae Baueraceae Bauhiniaceae Baxteriaceae BegoniaceaeBehniaceae B

elangeraceae Belloniaceae Bembiciaceae Berberidaceae Berberidopsidaceae Berryac

eaee BersamaceaeBertyaceae Berzeliaceae Besleriaceae Betaceae Betulaceae Biebe

rsteiniaceae Bifariaceae Bignoniaceae BischofiaceaeBixaceae Blakeaceae Blandfordia

ceae Blattiaceae Blepharocaryaceae Blitaceae Blyxaceae Boerlagellaceae Bolivaracea

eBombacaceae Bonnetiaceae Bontiaceae Boopidaceae Boraginaceae Borassaceae B

oroniaceae BoryaceaeBotryodendraceae Bougainvilleaceae Boweniaceae Brassicacea

e Brachycaulaceae Bretschneideraceae BrexiaceaeBromeliaceae Brownlowiaceae Br

unelliaceae Bruniaceae Brunoniaceae Brunsvigiaceae Bryoniaceae BuchneraceaeBuci

daceae Buddlejaceae Buglossaceae Bulbocodiaceae Bumeliaceae Bupleuraceae Burc

hardiaceae BurmanniaceaeBurseraceae Butneriaceae Butomaceae Buxaceae Byblida

ceae Byttneriaceae

Cabombaceae Cacaoaceae Cactaceae Caesalpiniaceae Caladiaceae Calamaceae Calce

olariaceae CalectasiaceaeCalendulaceae Callaceae Callicomaceae Calligonaceae Calli

traceae Callitrichaceae Calochortaceae CalophyllaceaeCalthaceae Calycanthaceae Ca

lyceraceae Cambogiaceae Camelliaceae Campanulaceae CampynemataceaeCanacom

yricaceae Canellaceae Cannabaceae Cannaceae Canopodaceae Canotiaceae Cansjera

ceae CapparaceaeCaprariaceae Caprifoliaceae Cardiopteridaceae Carduaceae Carica

ceae Carissaceae Carlemanniaceae CarpinaceaeCarpodetaceae Cartonemataceae Ca

ryocaraceae Caryophyllaceae Casearieaceae Cassiaceae CassipoureaceaeCassythacea

e Castaneaceae Castelaceae Casuarinaceae Catesbaeaceae Cathedraceae Cecropiace

ae CedraceaeCedrelaceae Celastraceae Celosiaceae Celtidaceae Centaureaceae Cen

trolepidaceae Centroplacaceae CepaceaeCephalanthaceae Cephalotaceae Cephalota

xaceae Cerastiaceae Ceratoniaceae Ceratophyllaceae CerberaceaeCercidiphyllaceae

Cercocarpaceae Cercodiaceae Cereaceae Cerinthaceae Ceroxylaceae Cestraceae Cev

alliaceaeChailletiaceae Chamaedoraceae Chamaemoraceae Chamelauciaceae Chaun

ochitonaceae Chelidoniaceae ChelonaceaeChenopodiaceae Chimonanthaceae Chingi

thamnaceae Chionographidaceae Chiranthodendraceae ChironiaceaeChlaenaceae Ch

loanthaceae Chloranthaceae Chloridaceae Chlorogalaceae Chrysobalanaceae Cicerac

eae CichoriaceaeCimicifugaceae Cinchoniaceae Circaeaceae Circaeasteraceae Cissac

eae Cistaceae Citraceae Clematidaceae CleomaceaeClethraceae Clusiaceae Cneorac

eae Cnestidaceae Cnicaceae Cobaeaceae Cochlospermaceae Cocosaceae Coffeaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Colchicaceae Coleogynaceae Columelliaceae Combretaceae Commelinaceae Comocl

adiaceae Compositae CompsoaceaeConantheraceae Connaraceae Conostylidaceae C

onvallariaceae Convolvulaceae Cordiaceae CoreopsidaceaeCoriandraceae Coriariacea

e Coridaceae Corispermaceae Cornaceae Corokiaceae Coronillaceae Correaceae Cor

rigiolaceaeCorsiaceae Corydalaceae Corylaceae Corynocarpaceae Coryphaceae Cost

aceae Cotyledonaceae CoulaceaeCoutariaceae Coutoubiaceae Crassulaceae Crescen

tiaceae Cressaceae Crinaceae Crocaceae CroomiaceaeCrossosomataceae Crotonacea

e Cruciferae Cryptaceae Crypteroniaceae Cryptocorynaceae CryptomeriaceaeCtenol

ophonaceae Cucurbitaceae Cunninghamiaceae Cunoniaceae Cupressaceae Curcumac

eae Curtisiaceae CuscutaceaeCyananthaceae Cyanastraceae Cynanchaceae Cyanella

ceae Cycadaceae Cyclanthaceae CyclantheraceaeCyclocheilaceae Cydoniaceae Cymo

doceaceae Cynaraceae Cynocrambaceae Cynomoriaceae Cyperaceae CyphiaceaeCyp

hocarpaceae Cypripediaceae Cyrillaceae Cyrtandraceae Cyrtanthaceae Cytinaceae

Dacrycarpaceae Dactylanthaceae Damasoniaceae Daphnaceae Daphniphyllaceae Das

ypogonaceae DatiscaceaeDaturaceae Daucaceae Davidiaceae Davidsoniaceae Decais

neaceae Deeringiaceae Degeneriaceae DelphiniaceaeDendrophthoaceae Columelliac

eae Detariaceae Dialypetalanthaceae Dianellaceae Dianthaceae DiapensiaceaeDicha

petalaceae Dichondraceae Diclidantheraceae Dicrastylidiaceae Dictamnaceae Didiere

aceae DidymelaceaeDidymocarpaceae Diegodendraceae Diervillaceae Digitalidaceae

e Dilatridaceae Dilleniaceae Dioaceae DionaeaceaeDioncophyllaceae Dioscoreaceae

Diosmaceae Diospyraceae Dipentodontaceae Diphylleiaceae Diplolaenaceae Dipsaca

ceaeDipterocarpaceae Dirachmaceae Disanthaceae Diselmaceae Dodonaeaceae Do

mbeyaceae Donatiaceae DorsteniaceaeDortmannaceae Doryanthaceae Drabaceae D

racaenaceae Dracontiaceae Drimyidaceae Droseraceae DrosophyllaceaeDryadaceae

Duabangaceae Duckeodendraceae Dulongiaceae Durantaceae Durionaceae Dysphani

aceae

Ebenaceae Eccremidaceae Ecdeiocoleaceae Echiaceae Echinopaceae Ehretiaceae Ela

eagnaceae ElaeocarpaceaeElegiaceae Elatinaceae Ellisiaceae Ellisiophyllaceae Elodea

ceae Elytranthaceae Embeliaceae Emblingiaceae EmmotaceaeEmpetraceae Encepha

lartaceae Engelhardtiaceae Enhalaceae Epacridaceae Ephedraceae Ephemeranthacea

eEpilobiaceae Epimediaceae Eragrostidaceae Eremolepidaceae Eremosynaceae Erica

ceae Erinaceae EriocaulaceaeEriogonaceae Eriospermaceae Erodiaceae Erycibaceae

Eryngiaceae Erysimaceae Erythroniaceae ErythropalaceaeErythrospermaceae Erythro

xylaceae Escalloniaceae Eschscholziaceae Eucomidaceae Eucommiaceae Eucryphiace

aeEuonymaceae Eupatoriaceae Euphorbiaceae Euphrasiaceae Euphroniaceae Eupo

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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matiaceae Eupteleaceae EuryalaceaeEustrephaceae Euthemidaceae Eutocaceae Exb

ucklandiaceae Exocarpaceae

Fabaceae Fagaceae Falcatifoliaceae Ferulaceae Festucaceae Ficaceae Ficoidaceae Fit

zroyaceae FlacourtiaceaeFlagellariaceae Flindersiaceae Foetidiaceae Forestieraceae

Fothergillaceae Fouquieriaceae Fragariaceae FrancoaceaeFrangulaceae Frankeniacea

e Fraxinaceae Fraxinellaceae Freycinetiaceae Fritillariaceae Fuchsiaceae Fumariacea

eFunkiaceae

Gaiadendraceae Galacaceae Galanthaceae Galaxiaceae Galedupaceae Galeniaceae G

aliaceae Garciniaceae GardeniaceaeGardneriaceae Garryaceae Geissolomataceae Ge

itonoplesiaceae Gelsemiaceae Geniostomaceae GentianaceaeGeonomataceae Geosir

idaceae Geraniaceae Gerrardinaceae Gesneriaceae Gethyllidaceae Gilliesiaceae Gina

lloaceaeGinkgoaceae Gisekiaceae Gladiolaceae Glaucidiaceae Glechomaceae Glinace

ae Globulariaceae Gnaphaliaceae GnetaceaeGoetzeaceae Gomortegaceae Gomphiac

eae Gomphrenaceae Gonystylaceae Goodeniaceae Gordoniaceae GouaniaceaeGoup

iaceae Gramineae Gratiolaceae Grewiaceae Greyiaceae Grielaceae Griseliniaceae Gr

onoviaceae GrossulariaceaeGrubbiaceae Guaiacanaceae Guamatelaceae Guettardac

eae Gunneraceae Gustaviaceae Guttiferae GyrocarpaceaeGyrostemonaceae

Hachetteaceae Haemanthaceae Haemodoraceae Halesiaceae Hallieraceae Halocarpa

ceae Halophilaceae HalophytaceaeHaloragaceae Hamamelidaceae Hanguanaceae H

aptanthaceae Harmandaceae Hebenstretiaceae HectorellaceaeHederaceae Hedyosm

aceae Hedyotidaceae Hedysaraceae Heisteriaceae Heleniaceae Heliamphoraceae H

elianthaceaeHelianthemaceae Helichrysaceae Heliconiaceae Helicteraceae Heliotropi

aceae Helleboraceae Heloniadaceae HelosidaceaeHelwingiaceae Hemerocallidaceae

Hemimeridaceae Hemiphylacaceae Henriqueziaceae Henslowiaceae Hermanniaceae

Hernandiaceae Herniariaceae Herreriaceae Hesperocallidaceae Heterantheraceae He

teropyxidaceae HeterostylaceaeHibbertiaceae Hibiscaceae Hilleriaceae Himantandra

ceae Hippocastanaceae Hippocrateaceae HippomanaceaeHippophaeaceae Hippurida

ceae Hirtellaceae Holacanthaceae Homaliaceae Hopkinsiaceae Hoplestigmataceae H

ordeaceaeHornschuchiaceae Hortensiaceae Hortoniaceae Hostaceae Houstoniaceae

Huaceae Huerteaceae HugoniaceaeHumbertiaceae Humiriaceae Hyacinthaceae Hyd

atellaceae Hydnoraceae Hydrangeaceae Hydrastidaceae HydrillaceaeHydrocharitacea

e Hydrocotylaceae Hydrogetonaceae Hydroleaceae Hydropeltidaceae Hydrophylacac

eae HydrophyllaceaeHydrostachyaceae Hymenocardiaceae Hypecoaceae Hyperanth

eraceae Hypericaceae Hypopityaceae HypoxidaceaeHypseocharitaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta

ceae ImperatoriaceaeInocarpaceae Inulaceae Ionidiaceae Iriarteaceae Iridaceae Irvi

ngiaceae Isnardiaceae Isophysidaceae IteaceaeStrasburgeriaceae Ixiaceae Ixioliriace

ae Ixonanthaceae

Jalapaceae Jamboliferaceae Japonoliriaceae Jasionaceae Jasminiaceae Johnsoniaceae

Joinvilleaceae JuglandaceaeJulianiaceae Juncaceae Juncaginaceae Juniperaceae Jus

siaeaceae Justiciaceae

Kadsuraceae Kaliphoraceae Kalmiaceae Kaniaceae Kiggelariaceae Kingdoniaceae Kin

giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae

Krameriaceae

Labiatae Lacandoniaceae Lachenaliaceae Lacistemataceae Lactoridaceae Lactucaceae

Lagerstroemiaceaee LamiaceaeLampsanaceae Lanariaceae Langsdorffiaceae Lantan

aceae Lapageriaceae Lardizabalaceae Lasiopetalaceae LauraceaeLawsoniaceae Laxm

anniaceae Lecythidaceae Ledaceae Ledocarpaceae Leeaceae Leguminosae Leitneriac

eae LemnaceaeLennoaceae Lentibulariaceae Lentiscaceae Leoniaceae Leonticaceae

Lepidobotryaceae LepidocarpaceaeLepidocaryaceae Lepidocerataceae Leptaulaceae

Leptospermaceae Lepturaceae Lepuropetalaceae LeuctenbergiaceaeLibocedraceae

Licaniaceae Ligustraceae Lilaeaceae Liliaceae Limeaceae Limnanthaceae Limnocharit

aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace

ae Linderniaceae Lindleyaceae Linnaeaceae LippayaceaeLiriaceae Liriodendraceae Li

ssocarpaceae Littorellaceae Loasaceae Lobeliaceaee Loganiaceae LomandraceaeLoni

ceraceae Lopeziaceae Lophiocarpaceae Lophiolaceae Lophiraceae Lophophytaceae L

ophopyxidaceae LoranthaceaeLotaceae Lowiaceae Lupulaceaee Luxemburgiaceae Lu

zuriagaceae Lychnidaceae Lyciaceae LyginiaceaeLygodisodeaceae Lysimachiaceae Ly

thraceae

Macarisiaceae Mackinlayaceae Madiaceae Maesaceae Magnoliaceae Malaceae Male

sherbiaceae MalortieaceaeMalpighiaceae Malvaceae Mangiaceae Manicariaceae Ma

paniaceae Marantaceae Marathraceae MarcgraviaceaeMartyniaceae Mastixiaceae

Matricariaceae Maundiaceae Mayacaceae Medeolaceae Medusagynaceae Peridiscac

eaeMelaleucaceae Melampyraceae Melanophyllaceae Melanthiaceae Melastomatac

eae Meliaceae Melianthaceae MelicaceaeMeliosmaceae Melittidaceae Melochiacea

e Memecylaceae Mendonciaceae Menispermaceae Menthaceae MenyanthaceaeMe

nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

Nypaceae Nyssaceae

Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

ceae Papilionaceae Pappophoraceae PapyraceaeParacryphiaceae Parasitaxaceae Par

ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

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ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 26: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

26

plants and landmarks are not as fixed as one might like trying to work out the relative

positions of parts is a good way of understanding morphology so long as one does not

try to out-do Procrustes Intermediates include intermediates found on the same

plant or on different plants Intermediates may observed during development or be

adult structures Thus one may find changes during development which suggest what

the nature of a particular structure is and very different stuctures on different plants

may turn out to be similar early in development Or one can simply compare different

taxa and see that structures that appear to be very different when comparing two

different taxa can be linked morphologically by looking at other taxa

Nevertheless the use of these criteria may not yield an unambiguous answer as to

what a structure is even given a solid phylogeny and an improved understanding of

development (see Jaramillo amp Kramer 2007 for a useful discussion) As Endress (2005c)

observed a number of features - position function development shape anatomy

histology gene activity and relationships to other taxa that clearly have petals - can be

used to distinguish a petal (for example) from other floral structures if a petal does not

have one of these features is it thereby not a petal Thus Maturen et al (2005) recently

found that floral organ diversity genes (B and C) were expressed in the large white

inflorescence bracts of Cornus (see also Costa et al 2005) Peney et al (2005) noted that

not all monosulcate pollen grains in monocots have the same developmental pathway

and that as a result such pollen might not have the same ancestral state Reeves and

Olmstead (2003) suggested that the genetic mechanisms causing monosymmetry in

Lamiales and Solanales were different and Serna and Martin (2006) described similar

problems with the development of hairs in Arabidopsis when compared with that of

hairs in Antirrhinum and Solanaceae Indeed as one perhaps might expect delimitation

of states and characters does not necessarily become easier with increasing knowledge

of development etc Thus Buzgo et al (2004) Matthews and Endress (2005) and others

have shown how hard it can be to distinguish between eg prophylls and other floral

structures as their behaviour is studied during the course of the development of the

flower Similarly at what concentration is a particular secondary metabolite deemed to

be present (Waterman (2007)

The validity of the approach used here that of fitting morphological variation to a

largely molecular-based tree may be questioned However I think it rather unlikely that

well-supported molecular branches will be overturned by morphological data Indeed

analyses of morphological data alone do provide support for many of the clades evident

in molecule-only analyses and analyses with morphological and molecular data

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

27

together may lead to increased support for clades (eg Hufford 1992 Nandi et al 1998

[but adding morphological data reduces support for a number of critical clades too]

Doyle amp Endress 2000 however in none of these papers is the use of morphology

without ambiguity) It is unfortunately clear that the use of morphology alone may not

suggest problems in the phylogenetic placement of taxa that later turn out to have been

wrongly included (eg Zhang et al 1992) Thus I have been wary of putting much weight

on clades that have only morphological support but note that I have treated molecule-

based clades with low bootstrap or jacknife support values (esp below 70) or low

posterior probabilies (below 095) likewise Although I may have been mistaken in

placing so much emphasis on molecular data in terms of providing the basic

phylogenetic framework for angiosperms morphological and molecular data are only

very rarely in irreconcilably strong conflict There are indeed a few places where the

conflict seems extreme These include the relative positions of the Monimiaceae and

Hernandiaceae (Laurales) the position of Hanguanaceae (Commelinales [as here] or

Zingiberales) and of Triplostegia (is it in Dipsacaceae or Valerianaceae - see

Dipsacales) Fossils are unlikely to affect the topologies of the trees presented here but

see below for their importance in understanding morphological evolution in general and

the evolution of angiosperms in particular Nevertheless some largely reject the idea

that trees based on molecular data alone can recover phylogenetic relationships

especially when branching points are old and prefer to used trees based on analysis of

morphological data including those taken from fossils (Hilton amp Bateman 2006 Farjon

2007)

All in all however the extent of the congruence between morphological and

molecular data is impressive and heartening and many clades can be characterised

morphologically It seemed in 1998 that there were no unambiguous morphological

synapomorphies for angiosperm orders (K Bremer 2000) and this is still true if by

unambiguous is meant non-homoplasious However many orders and other clades

have synapomorphies even if these may be indistinguishable at least at the current

level of morphological and developmental knowledge from parallel occurences

elsewhere As our morphological knowledge increases so too does the number of

apomorphies

Endress and Matthews (2006a) emphasize the importance of tendencies and

developmental constraints when thinking about characters of clades within the rosids -

many characters or character combinations occur in a rather sporadic fashion within

the clade and apparently notably less frequently outside it There are a number of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

28

examples of tendencies here and in the individual order pages such as the features

enclosed by parentheses in groups above the level of family eg see (cuticular waxes

as aggregated rodlets) for the commelinids and the discussion of the distribution of

polyandry that is flowers with numerous stamens in the asterid I + II groups etc

Tendencies may involve more than single characters When thinking of the

associations of features like integument thickness and vascularization of the integument

(Endress amp Matthews 2006a) one can imagine fairly simple developmental

preconditions being operative As Endress and Matthews (2006a) note it is difficult to

imagine vascular tissue developing in an integument only two cells thick Changes here

would be loosely correlated if morphologically linked However the strongly correlated

changes noted by Givnish et al (2005) are ecologically linked but are presumably

morphologicallydevelopmentally independent When there seem to be characters

evolving more or less together tests can be carried out to see if the changes are

concentrated on certain branches of the tree (eg Maddison 1990 Sanderson 1991

Maddison amp Maddison 2000) Such tests have rarely been carried out at higher levels in

angiosperms

One of the most striking examples of a tendency is the distribution of N-fixation

restricted as it is to a monophyletic group of four clades although it has arisen seven (or

perhaps several more) times independently within the N-fixing clade and several

members of two quite different kinds of bacteria are involved (eg Clawson et al 2004

Elliott et al 2007 Sprent amp James 2007) Other examples of work that bears on the issue

of tendencies include the findings that flowers of polysymmetrical Arabidopsis have

genes like TCP1 that are expressed asymmetrically during early development and TCP1

is a probable orthologue of the well-known CYC gene of Antirrhinum that is involved in

the development of monosymmetric slowers there (Cubas et al 2001 Costa et al 2005

etc) Parallelism might build on this underlying morphologically cryptic monosymmetry

even if details of the genetic mechanisms causing the monosymmetry evident in

particular groups may be different (Reeves amp Olmstead 2003 Cubas 2004 see above)

Similarly there have been several recent suggestions that the capability to synthesise a

particular metabolite may be switched off but not lost and so can sometimes be

reacquired (eg Wink amp Witte 1983 Wink 2003 Liscombe et al 2005 Larsson 2007

Waterman 2007) Hence perhaps the rather spotty distribution of many secondary

metabolites like ellagic acid the indole alkaloid camptothecin iridoids etc when

considered in the context of phylogenies As a non-botanical example - but a rather nice

one - Salwini-Plawen and Mayr (1961) suggested some time ago that there has been

considerable parallelism (40-65 or more independent origins) in the evolution of eyes in

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

29

metazoans However Pax 6 seems to be a master control gene that is involved in eye

formation perhaps throughout all bilateralians (eg Gehring amp Ikeo 1999 Erwin amp

Davidson 2002) which changes the nature of the problem Note however that in

general our invocation of evolutionary tendencies to explain the patterns of

distributions of characters is really an admission that we do not understand those

patterns

As we find out more about variation we will probably find fewer and fewer features

constant throughout a group Most unqualified statements of presence and absence

should properly be qualified as usually present or usually absent if one is thinking of

the characterisations on the order pages as encompassing the total variation within a

clade Thus Pistia alone among monocots as so far known has sieve tube plastids with

starch grains not protein crystals However this does not affect the fact that sieve tube

plastids with cuneate protein crystals and lacking starch grains are an apomorphy for

monocots Furthermore fossils in a number of cases suggest character combinations

unknown in extant taxa as may be seen in the discussions of Fagaceae Platanaceae

Iteaceae Calycanthaceae etc Confusing the issue there may be questions as to where

exactly on the tree a particular fossil is to be placed (eg see Nymphaeaceae

Calycanthaceae Archaefructus etc)

To summarize given our current understandings of both phylogenies and

characters evolution of some characters in which we are interested seems very labile

(see eg D Soltis et al 2005b Endress and Matthews 2006a Stevens 2006b Ekman et

al 2008 for a good recent study) and I have been cautious when talking about character

evolution Much effort must continue to be spent in summarizing characters of well-

established clades at all levels providing features by which they may be recognized and

signaling synapomorphies Remember that (1) the basic morphological anatomical and

chemical knowledge of many critical taxa is woefully incomplete (2) different

assumptions about character evolution may greatly affect the position of

synapomorphies on trees (3) in many cases relationships within and between many

groups are too uncertain at present to worry very much about synapomorphies and

(4) we must be clear about what we do and do not not know As mentioned above I

have indicated in a number of places where there are particularly important

uncertainties as to where characters should be placed on the tree Nevertheless it is a

relatively easy matter to update notes such as these and it can be a simple matter to

incorporate new data on characters that have never before been considered in the

context of a tree Much basic - and unfortunately perhaps unfashionable - work must be

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

30

carried to clarify the distribution of morphological anatomical and chemical characters

for examples of what can to be done see the work of P K Endress and collaborators

(floral morphology and development) and S R Jensen and collaborators (iridoids) But

acquiring information about nodal anatomy cell and tissue distribution nodal anatomy

and the like is not difficult at all and should be generally encouraged

But all this is merely a necesary prelude to the understanding of evolution For this

one needs to know a lot more including dating the phylogeny understanding the

function(s) of characters factoring in relevant aspects of the palaeoenvironment in

which an apomorphy first appeared etc

SUMMARY OF THE SYSTEM

Below is a formalised summary of the relationships within orders of the families of seed

plants There are a few families that are not recognised even as options in APG II and

vice versa as well as a few extra orders The families to be recognised in the most

recent edition of Mabberleys The Plant Book (Mabberley 2008) are also largely

consistent with those below But all differences are trivial and will - I hope - eventually

disappear

Square brackets - [] - enclose clades the plus sign - + - designates sister taxa a

comma - - denotes part of a polytomy and quotation marks - - denotes a

paraphyletic group

SEED PLANTS

GYMNOSPERMS

Cycadales

Cycadaceae + Zamiaceae

Ginkgoales

Ginkgoaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

31

Pinales

Pinaceae [[Araucariaceae [Podocarpaceae + Phyllocladaceae]] [Sciadopityaceae

[Taxaceae + Cupressaceae]]]

Gnetales

Ephedraceae [Gnetaceae + Welwitschiaceae]

ANGIOSPERMSFLOWERING PLANTS

Amborellales

Amborellaceae

Nymphaeales

Hydatellaceae [Cabombaceae + Nymphaeaceae]

Austrobaileyales

Austrobaileyaceae [Schisandraceae + Trimeniaceae]

Chloranthales

Chloranthaceae

MAGNOLIIDS

Magnoliales

Myristicaceae [Magnoliaceae [[Himantandraceae + Degeneriaceae] [Eupomatiaceae +

Annonaceae]]]

Laurales

Calycanthaceae [[Siparunaceae [Gomortegaceae + Atherospermataceae]] [Monimiaceae

[Hernandiaceae + Lauraceae]]]

Canellales

Canellaceae + Winteraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

32

Piperales

[Hydnoraceae + Aristolochiaceae] [Piperaceae + Saururaceae]

MONOCOTS

Acorales

Acoraceae

Alismatales

Araceae [Tofieldiaceae [[Alismataceae [Hydrocharitaceae + Butomaceae]]

[Scheuchzeriaceae [Aponogetonaceae [Juncaginaceae [Maundiaceae [[Posidoniaceae

[Ruppiaceae + Cymodoceaceae]] [Zosteraceae + Potamogetonaceae]]]]]]]

Petrosaviales

Petrosaviaceae

Dioscoreales

Nartheciaceae [[Taccaceae + Thismiaceae] [Burmanniaceae + Dioscoreaceae]]

Pandanales

Velloziaceae Triuridaceae Stemonaceae [Pandanaceae + Cyclanthaceae]

Liliales

Corsiaceae [Campynemataceae [Petermanniaceae [Colchicaceae + Alstroemeriaceae]]

Melanthiaceae [[Philesiaceae + Rhipogonaceae] [Smilacaceae + Liliaceae]]]

Asparagales

Orchidaceae [[Boryaceae [Blandfordiaceae [Lanariaceae [Asteliaceae + Hypoxidaceae]]]]

[[Ixioliriaceae + Tecophilaeaceae] [Doryanthaceae [Iridaceae [Xeronemataceae

[Xanthorrhoeaceae [Alliaceae + Asparagaceae]]]]]]]

COMMELINIDS

Unplaced

Dasypogonaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

33

Arecales

Arecaceae

Poales

[Typhaceae + Bromeliaceae] [Rapateaceae [[Thurniaceae [Juncaceae + Cyperaceae]]

[[Mayacaceae [Eriocaulaceae + Xyridaceae]] [Flagellariaceae [Anarthriaceae

[Centrolepidaceae + Restionaceae]]] [Joinvilleaceae [Ecdeiocoleaceae + Poaceae]]]]]]]

Commelinales

[Commelinaceae + Hanguanaceae] [Philydraceae [Haemodoraceae + Pontederiaceae]]

Zingiberales

Musaceae [Strelitziaceae + Lowiaceae] Heliconiaceae [[Cannaceae + Marantaceae]

[Costaceae + Zingiberaceae]]

Ceratophyllales

Ceratophyllaceae

EUDICOTS

Ranunculales

Eupteleaceae [Papaveraceae [[[Lardizabalaceae + Circaeasteraceae] [Menispermaceae

[Berberidaceae + Ranunculaceae]]]]

Sabiales

Sabiaceae

Proteales

Nelumbonaceae [Platanaceae + Proteaceae]

Trochodendrales

Trochodendraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

34

Buxales

Haptanthaceae [Buxaceae + Didymelaceae]

CORE EUDICOTS

Gunnerales

Gunneraceae + Myrothamnaceae

Dilleniales

Dilleniaceae

Saxifragales

Peridiscaceae [[Paeoniaceae [Altingiaceae [Hamamelidaceae [Cercidiphyllaceae +

Daphniphyllaceae]]]] [[Crassulaceae [Aphanopetalaceae [Tetracarpaeaceae

[Penthoraceae + Haloragaceae]]]] [Iteaceae [Grossulariaceae + Saxifragaceae]]]]

Cynomoriaceae unplaced

Vitales

Vitaceae

ROSIDS

FABIDROSID I

Zygophyllales

Krameriaceae + Zygophyllaceae

Celastrales

Lepidobotryaceae + Celastraceae

Oxalidales

Huaceae [[Connaraceae + Oxalidaceae] [Cunoniaceae [Elaeocarpaceae [Brunelliaceae +

Cephalotaceae]]]]

Malpighiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

35

[Achariaceae [Goupiaceae [Violaceae + Passifloraceae] [Lacistemataceae Salicaceae]]

[Lophopyxidaceae Putranjivaceae] [Ctenolophonaceae [Erythroxylaceae +

Rhizophoraceae]] Linaceae Ixonanthaceae Humiriaceae Irvingiaceae

Centroplacaceae Pandaceae [[Bonnetiaceae + Clusiaceae] [Calophyllaceae

[Hypericaceae + Podostemaceae]]] [Malpighiaceae + Elatinaceae] Ochnaceae

[Peraceae [Rafflesiaceae + Euphorbiaceae]] [Phyllanthaceae + Picrodendraceae]

[Balanopaceae [[Trigoniaceae + Dichapetalaceae] [Chrysobalanaceae +

Euphroniaceae]]] Caryocaraceae

N-FIXING CLADE

Fabales

Quillajaceae [Fabaceae [Polygalaceae + Surianaceae]]

Rosales

Rosaceae [[Barbeyaceae [Elaeagnaceae Dirachmaceae Rhamnaceae]] [Ulmaceae

[Cannabaceae [Moraceae + Urticaceae]]]]

Cucurbitales

Anisophylleaceae [[Corynocarpaceae + Coriariaceae] [Cucurbitaceae [Tetramelaceae

[Datiscaceae + Begoniaceae]]]] Apodanthaceae

Fagales

Nothofagaceae [Fagaceae [[Myricaceae + Juglandaceae] [Casuarinaceae

[Ticodendraceae + Betulaceae]]]]

MALVIDROSID II

pgtGeraniales

Geraniaceae [[Melianthaceae + Francoaceae] [Vivianaceae + Ledocarpaceae]]

Myrtales

Combretaceae [[Onagraceae + Lythraceae] [[Vochysiaceae + Myrtaceae]

[Melastomataceae [Crypteroniaceae [Alzateaceae + Penaeaceae]]]]]

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

36

Crossosomatales

[Staphyleaceae [Guamatelaceae [Crossosomataceae + Stachyuraceae]]] [Aphloiaceae

[Geissolomataceae + Strasburgeriaceae]]

Picramniales

Picramniaceae

Sapindales

Biebersteiniaceae Nitrariaceae [Kirkiaceae [Anacardiaceae + Burseraceae]]

[Sapindaceae [Simaroubaceae Rutaceae Meliaceae]]

Huerteales

Gerradinaceae [Dipentodontaceae + Tapisciaceae]

Malvales

Neuradaceae [Thymelaeaceae [Sphaerosepalaceae Bixaceae [Cistaceae

[Sarcolaenaceae + Dipterocarpaceae]] [Cytinaceae + Muntingiaceae] Malvaceae]]

Brassicales

[Akaniaceae + Tropaeolaceae] [[Moringaceae + Caricaceae] [Setchellanthaceae

[Limnanthaceae [[Koeberliniaceae [Bataceae + Salvadoraceae]] [Emblingiaceae

[Pentadiplandraceae [Gyrostemonaceae Resedaceae Stixis and relatives] Tovariaceae

[Capparaceae [Cleomaceae + Brassicaceae]]]]]]]]

Berberidopsidales

Aextoxicaceae + Berberidopsidaceae

Santalales

Erythropalaceae Olacaceae [[Loranthaceae [Misodendraceae + Schoepfiaceae]]

[Opiliaceae + Santalaceae]] Balanophoraceae unplaced

Caryophyllales

[[Droseraceae [Nepenthaceae [Drosophyllaceae [Ancistrocladaceae +

Dioncophylleaceae]]]]] [[Frankeniaceae + Tamaricaceae] [Polygonaceae +

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

37

Plumbaginaceae]]] [Rhabdodendraceae [Simmondsiaceae [[Asteropeiaceae +

Physenaceae] [[Caryophyllaceae [Achatocarpaceae + Amaranthaceae]]

[Stegnospermataceae [Limeaceae [[Lophocarpaceae [Barbeuiaceae [Aizoaceae

[Sarcobataceae Phytolaccaceae Nyctaginaceae]]]] [Molluginaceae [Halophytaceae

Montiaceae Basellaceae Didiereaceae [Talinaceae [Portulacaceae [Anacampseros etc +

Cactaceae]]]]]]]]]]]]]

ASTERIDS

Cornales

[Cornaceae + Nyssaceae] [Hydrangeaceae + Loasaceae] Hydrostachyaceae [Curtisiaceae

+ Grubbiaceae]

Ericales

[Balsaminaceae [Marcgraviaceae + Tetrameristaceae]] [[Polemoniaceae +

Fouquieraceae] Lecythidaceae [[Sladeniaceae + Pentaphylacaceae] [Sapotaceae

[Ebenaceae [Maesaceae [Theophrastaceae [Myrsinaceae + Primulaceae]]]]]

[Mitrastemonaceae Theaceae [Symplocaceae [Styracaceae + Diapensiaceae]]

[[Sarraceniaceae [Roridulaceae + Actinidiaceae] [Clethraceae [Cyrillaceae +

Ericaceae]]]]]]

LAMIIDASTERID I

Unplaced

Metteniusaceae Oncothecaceae Icacinaceae and some unplaced ex-Icacinaceae

Garryales

Garryaceae + Eucommiaceae

Unplaced

Boraginaceae Vahliaceae

Gentianales

Rubiaceae [Gentianaceae [Loganiaceae [Gelsemiaceae + Apocynaceae]]]

Lamiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

38

Plocospermataceae [Oleaceae [Tetrachondraceae [Calceolariaceae

[Gesneriaceae Peltanthera [Pedaliaceae Carlemanniaceae Martyniaceae Stilbaceae

Plantaginaceae Linderniaceae Bignoniaceae Verbenaceae Lamiaceae Paulowniaceae

Schlegeliaceae Thomandersiaceae Phrymaceae Rehmannia Acanthaceae

Scrophulariaceae Orobanchaceae Byblidaceae Lentibulariaceae]]]]]

Solanales

[Montiniaceae [Sphenocleaceae + Hydroleaceae]] [Convolvulaceae + Solanaceae]

CAMPANULIDASTERID II

Aquifoliales

[[Cardiopteridaceae + Stemonuraceae] [Aquifoliaceae [Helwingiaceae +

Phyllonomaceae]]

Asterales

[Rousseaceae + Campanulaceae] Pentaphragmataceae [[Alseuosmiaceae [Phellinaceae

+ Argophyllaceae]] Stylidiaceae [Menyanthaceae [Goodeniaceae [Calyceraceae +

Asteraceae]]]]

Escalloniales

Escalloniaceae

Bruniales

[Bruniaceae + Columelliaceae]

Apiales

Pennantiaceae [Torricelliaceae [Griseliniaceae [Pittosporaceae [Araliaceae

[Myodocarpaceae + Apiaceae]]]]]

Paracryphiales

Paracryphiaceae

Dipsacales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

39

Adoxaceae [Diervillaceae [Caprifoliaceae [Linnaeaceae [Morinaceae [Dipsacaceae +

Valerianaceae]]]]]

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MAIN GROUPINGS

Asterids commelinids core eudicots asterid 1 asterid

2 eudicots gymnosperms Magnoliophyta monocots N-fixing claderosids seed plants

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Abietales Acanthales Acerales Acorales Actinidiales Actinostrobales Adoxales Aescul

ales Agavales Aizoales AkanialesAlismatales Alliales Alseuosmiales Alstroemeriales

Altingiales Amaranthales Amaryllidales Amborellales AmbrosialesAmmiales Amomal

es Ancistrocladales Anisophylleales Annonales Anthobolales Apiales Apocynales Apo

nogetonalesAquifoliales Arales Araliales Aralidiales Araucariales Arecales Aristolochi

ales Athrotaxidales Asarales AsclepiadalesAsparagales Asphodelales Asparagales Ast

eliales Atriplicales Aucubales Austrobaileyales Avenales

Balanitales Balanopales Balanophorales Balsaminales Barbeyales Barclayales Batales

Begoniales BerberidalesBerberidopsidales Betulales Biebersteiniales Bignoniales Bix

ales Boraginales Brassicales Brexiales Bromeliales BrunialesBrunoniales Burmannial

es Burserales Butomales Buxales Byblidales

Cactales Callitrichales Calycanthales Calycerales Campanulales Campynematales Can

ellales Cannales CapparalesCaprifoliales Cardiopteridales Carduales Caricales Carle

manniales Caryophyllales Cassiales Casuarinales CelastralesCentrolepidales Cephalot

ales Cephalotaxales Ceratophyllales Cercidiphyllales Chenopodiales Chironiales Chlor

anthalesChrysobalanales Cinchonales Circaeasterales Cistales Citrales Cocosales Col

chicales Columelliales CombretalesCommelinales Connarales Convolvulales Coriarial

es Cornales Corylales Corynocarpales Crassulales CrossosomatalesCucurbitales Cunn

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

40

inghamiales Cunoniales Cupressales Cycadales Cyclanthales Cymodoceales Cynarales

CynomorialesCyperales Cyrillales Cytinales

Daphnales Daphniphyllales Dasypogonales Datiscales Degeneriales Desfontainiales D

iapensiales Didymelales DillenialesDioales Dioncophyllales Dioscoreales Diospyrales

Dipentodontales Dipsacales Droserales

Ebenales Echiales Elaeagnales Elaeocarpales Elatinales Elodeales Emmotales Empetr

ales Ephedrales Ericales EriocaulalesErythropalaless Escalloniales Eucommiales Euph

orbiales Eupomatiales Eupteleales Euryalaless

Fabales Fagales Falcatifoliales Ficales Flacourtiales Flagellariales Fouquieriales Franc

oales Frangulales

Galiales Garryales Geissolomatales Gentianales Geraniales Gesneriales Ginkgoales G

laucidiales Globulariales GnetalesGoodeniales Greyiales Griseliniales Grossulariales

Grubbiales Gunnerales Gyrocarpales Gyrostemonales

Haemodorales Haloragales Hamamelidales Hanguanales Heisteriales Helleborales He

lwingiales Himantandrales HippuridalesHomaliales Hortensiales Huales Huerteales

Hydatellales Hydnorales Hydrangeales Hydrastidales HydrocharitalesHydropeltidales

Hydrostachyales Hypericales Hypoxidales

Icacinales Illiciales Iridales Irvingales Iteales Ixiales

Jasminales Juglandales Julianiales Juncaginales Juncales

Lacistematales Lactoridales Lamiales Lardizabalales Laurales Lecythidales Ledocarpal

es Leitneriales LentibularialesLigustrales Liliales Limnanthales Linales Liriales Loasal

es Lobeliales Loganiales Lonicerales Loranthales LowialesLythrales

Magnoliales Malpighiales Malvales Marathrales Marcgraviales Mayacales Medusagy

nales Medusandrales MelanthialesMelastomatales Meliales Melianthales Meliosmal

es Menispermales Menyanthales Metteniusales MitrastemonalesMiyoshiales Moni

miales Moringales Musales Myricales Myristicales Myrothamnales Myrsinales Myrta

les

Najadales Nandinales Narcissales Nartheciales Nelumbonales Nepenthales Neuradal

es Nitrariales Nolanales NothofagalesNyctaginales Nymphaeales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Ochnales Oenotherales Olacales Oleales Onagrales Oncothecales Opuntiales Orchid

ales Oxalidales

Saxifragales Pandales Pandanales Papaverales Paracryphiales Parasitaxales Paridales

Parnassiales PassifloralesPenaeales Pennantiales Pentaphragmatales Peridiscales P

etiveriales Petrosaviales Phellinales Philydrales PhyllanthalesPhysenales Phytolaccal

es Picramniales Pinales Pinguiculales Piperales Pittosporales Plantaginales Platanale

sPlumbaginales Poales Podocarpales Podophyllales Podostemales Polemoniales Pol

ygalales Polygonales PontederialesPortulacales Posidoniales Potamogetonales Primu

lales Proteales

Quercales Quillajales Quintiniales

Rafflesiales Ranunculales Rapateales Resedales Restionales Rhabdodendrales Rhamn

ales Rhinanthales RhizophoralesRhodorales Rhoipteleales Roridulales Rosales Rouss

eales Rubiales Ruppiales Rutales

Sabiales Salicales Salvadorales Samolales Samydales Sanguisorbales Santalales Sapin

dales Sapotales SarracenialesSaxegotheales Saxifragales Scheuchzeriales Sciadopitya

les Scleranthales Scrophulariales Scyphostegiales SedalesSilenales Simmondsiales S

milacales Solanales Sphenocleales Sphenostemonales Stangeriales Stellariales Stemo

nalesStilbales Stylidiales Styracales Surianales

Taccales Tamales Tamaricales Taxales Taxodiales Tecophilaeales Terebinthales Tern

stroemiales Theales TheligonalesThymelaeales Tiliales Tofieldiales Torricelliales Tov

ariales Tribelales Trilliales Trimeniales Triuridales TrochodendralesTropaeolales Turn

erales Typhales

Ulmales Urticales

Vacciniales Vahliales Vallisneriales Velloziales Veratrales Verbenales Viburnales Vinc

ales Violales Viscales VitalesVochysiales

Welwitschiales Winterales

Xanthorrhoeales Ximeniales Xyridales

Zamiales Zingiberales Zosterales Zygophyllales

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which the genus of the basionym is to be found

Abietaceae Abolbodaceae Abrophyllaceae Acalyphaceae Acanthaceae [Acanthochla

mydaceae] Acarnaceaee AceraceaeAchariaceae Achatocarpaceae Achradaceae Achy

ranthaceae Acmopylaceae Acoraceae Acristaceae ActaeaceaeActinidiaceae Actinota

ceae Amentotaxaceae Adenogrammaceae Adoxaceae Aegialitidaceae Aegicerataceae

Aegilopaceae Aeginetiaceaee Aegiphilaceae Aesculaceae Aextoxicaceae Agapanthac

eae Agavaceae AgdestidaceaeAgialidaceae Agrimoniaceae Agrostidaceae Ailanthace

ae Aitoniaceae Aizoaceae Akaniaceae Alangiaceae AlchemillaceaeAldrovandaceae A

lismataceae Alliaceae Allioniaceae Allophylaceae Aloaceae Alopecuraceae Alpiniacea

e AlseuosmiaceaeAlsinaceae Alsodeiaceae Alstroemeriaceae Altingiaceae Alzateacea

e Amaranthaceae Amaryllidaceae Amborellaceae Ambrosiaceae Amentotaxaceae A

mmanniaceae Ammiaceae Amomaceae Ampelidopsaceae Amygdalaceae Amyridacea

eAnacardiaceae Anagallidaceae Anarthriaceae Anchusaceae Ancistrocladaceae Andr

omedaceae AndropogonaceaeAndrostachyaceae Androsynaceae Anemarrhenaceae

Anemonaceae Angelicaceae Anisophylleaceae AnnonaceaeAnomochloaceae Anopter

aceae Anrederaceae Anthemidaceae Anthericaceae Anthobolaceae Antidesmataceae

Antirrhinaceae Antoniaceae Aparinaceae Aphanopetalaceae Aphloiaceae Aphyllanth

aceae Apiaceae ApocynaceaeApodanthaceae Aponogetonaceae Aporusaceae Apose

ridaceae Apostasiaceae Aptandraceae AquifoliaceaeAquilariaceae Aquilegiaceae A

raceae Aragoaceae Araliaceae Aralidiaceaee Araucariaceae Arbutaceae Arceuthidace

aeArceuthobiaceae Arctostaphylaceae Arctotidaceae Ardisiaceae Arecaceae Argoph

yllaceae Arisaraceae AristolochiaceaeAristoteliaceae Arjonaceae Armeriaceae Artem

isiaceae Arthrotaxidaceae Artocarpaceae Arundinaceae ArundinellaceaeAsaraceae A

sclepiadaceae Ascyraceaee Asiraceae Asparagaceae Asperulaceae Asphodelaceae As

pidistraceaeeAsteliaceae Asteraceae Asteranthaceae Asteropeiaceae Astragalaceae

Astrocarpaceae AthanasiaceaeAtherospermataceae Atriplicaceae Atropaceae Aucuba

ceae Aurantiaceae Austrobaileyaceae Austrotaxaceae AvenaceaeAverrhoaceae Avet

raceae Avicenniaceae Azaleaceae Azimaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Balanitaceae Balanopaceae Balanophoraceae Balsameaceae Balsaminaceae Bambusa

ceae Barbaceniaceae BarbeuiaceaeBarbeyaceae Barclayaceae Barringtoniaceae Base

llaceae Bataceae Baueraceae Bauhiniaceae Baxteriaceae BegoniaceaeBehniaceae B

elangeraceae Belloniaceae Bembiciaceae Berberidaceae Berberidopsidaceae Berryac

eaee BersamaceaeBertyaceae Berzeliaceae Besleriaceae Betaceae Betulaceae Biebe

rsteiniaceae Bifariaceae Bignoniaceae BischofiaceaeBixaceae Blakeaceae Blandfordia

ceae Blattiaceae Blepharocaryaceae Blitaceae Blyxaceae Boerlagellaceae Bolivaracea

eBombacaceae Bonnetiaceae Bontiaceae Boopidaceae Boraginaceae Borassaceae B

oroniaceae BoryaceaeBotryodendraceae Bougainvilleaceae Boweniaceae Brassicacea

e Brachycaulaceae Bretschneideraceae BrexiaceaeBromeliaceae Brownlowiaceae Br

unelliaceae Bruniaceae Brunoniaceae Brunsvigiaceae Bryoniaceae BuchneraceaeBuci

daceae Buddlejaceae Buglossaceae Bulbocodiaceae Bumeliaceae Bupleuraceae Burc

hardiaceae BurmanniaceaeBurseraceae Butneriaceae Butomaceae Buxaceae Byblida

ceae Byttneriaceae

Cabombaceae Cacaoaceae Cactaceae Caesalpiniaceae Caladiaceae Calamaceae Calce

olariaceae CalectasiaceaeCalendulaceae Callaceae Callicomaceae Calligonaceae Calli

traceae Callitrichaceae Calochortaceae CalophyllaceaeCalthaceae Calycanthaceae Ca

lyceraceae Cambogiaceae Camelliaceae Campanulaceae CampynemataceaeCanacom

yricaceae Canellaceae Cannabaceae Cannaceae Canopodaceae Canotiaceae Cansjera

ceae CapparaceaeCaprariaceae Caprifoliaceae Cardiopteridaceae Carduaceae Carica

ceae Carissaceae Carlemanniaceae CarpinaceaeCarpodetaceae Cartonemataceae Ca

ryocaraceae Caryophyllaceae Casearieaceae Cassiaceae CassipoureaceaeCassythacea

e Castaneaceae Castelaceae Casuarinaceae Catesbaeaceae Cathedraceae Cecropiace

ae CedraceaeCedrelaceae Celastraceae Celosiaceae Celtidaceae Centaureaceae Cen

trolepidaceae Centroplacaceae CepaceaeCephalanthaceae Cephalotaceae Cephalota

xaceae Cerastiaceae Ceratoniaceae Ceratophyllaceae CerberaceaeCercidiphyllaceae

Cercocarpaceae Cercodiaceae Cereaceae Cerinthaceae Ceroxylaceae Cestraceae Cev

alliaceaeChailletiaceae Chamaedoraceae Chamaemoraceae Chamelauciaceae Chaun

ochitonaceae Chelidoniaceae ChelonaceaeChenopodiaceae Chimonanthaceae Chingi

thamnaceae Chionographidaceae Chiranthodendraceae ChironiaceaeChlaenaceae Ch

loanthaceae Chloranthaceae Chloridaceae Chlorogalaceae Chrysobalanaceae Cicerac

eae CichoriaceaeCimicifugaceae Cinchoniaceae Circaeaceae Circaeasteraceae Cissac

eae Cistaceae Citraceae Clematidaceae CleomaceaeClethraceae Clusiaceae Cneorac

eae Cnestidaceae Cnicaceae Cobaeaceae Cochlospermaceae Cocosaceae Coffeaceae

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Colchicaceae Coleogynaceae Columelliaceae Combretaceae Commelinaceae Comocl

adiaceae Compositae CompsoaceaeConantheraceae Connaraceae Conostylidaceae C

onvallariaceae Convolvulaceae Cordiaceae CoreopsidaceaeCoriandraceae Coriariacea

e Coridaceae Corispermaceae Cornaceae Corokiaceae Coronillaceae Correaceae Cor

rigiolaceaeCorsiaceae Corydalaceae Corylaceae Corynocarpaceae Coryphaceae Cost

aceae Cotyledonaceae CoulaceaeCoutariaceae Coutoubiaceae Crassulaceae Crescen

tiaceae Cressaceae Crinaceae Crocaceae CroomiaceaeCrossosomataceae Crotonacea

e Cruciferae Cryptaceae Crypteroniaceae Cryptocorynaceae CryptomeriaceaeCtenol

ophonaceae Cucurbitaceae Cunninghamiaceae Cunoniaceae Cupressaceae Curcumac

eae Curtisiaceae CuscutaceaeCyananthaceae Cyanastraceae Cynanchaceae Cyanella

ceae Cycadaceae Cyclanthaceae CyclantheraceaeCyclocheilaceae Cydoniaceae Cymo

doceaceae Cynaraceae Cynocrambaceae Cynomoriaceae Cyperaceae CyphiaceaeCyp

hocarpaceae Cypripediaceae Cyrillaceae Cyrtandraceae Cyrtanthaceae Cytinaceae

Dacrycarpaceae Dactylanthaceae Damasoniaceae Daphnaceae Daphniphyllaceae Das

ypogonaceae DatiscaceaeDaturaceae Daucaceae Davidiaceae Davidsoniaceae Decais

neaceae Deeringiaceae Degeneriaceae DelphiniaceaeDendrophthoaceae Columelliac

eae Detariaceae Dialypetalanthaceae Dianellaceae Dianthaceae DiapensiaceaeDicha

petalaceae Dichondraceae Diclidantheraceae Dicrastylidiaceae Dictamnaceae Didiere

aceae DidymelaceaeDidymocarpaceae Diegodendraceae Diervillaceae Digitalidaceae

e Dilatridaceae Dilleniaceae Dioaceae DionaeaceaeDioncophyllaceae Dioscoreaceae

Diosmaceae Diospyraceae Dipentodontaceae Diphylleiaceae Diplolaenaceae Dipsaca

ceaeDipterocarpaceae Dirachmaceae Disanthaceae Diselmaceae Dodonaeaceae Do

mbeyaceae Donatiaceae DorsteniaceaeDortmannaceae Doryanthaceae Drabaceae D

racaenaceae Dracontiaceae Drimyidaceae Droseraceae DrosophyllaceaeDryadaceae

Duabangaceae Duckeodendraceae Dulongiaceae Durantaceae Durionaceae Dysphani

aceae

Ebenaceae Eccremidaceae Ecdeiocoleaceae Echiaceae Echinopaceae Ehretiaceae Ela

eagnaceae ElaeocarpaceaeElegiaceae Elatinaceae Ellisiaceae Ellisiophyllaceae Elodea

ceae Elytranthaceae Embeliaceae Emblingiaceae EmmotaceaeEmpetraceae Encepha

lartaceae Engelhardtiaceae Enhalaceae Epacridaceae Ephedraceae Ephemeranthacea

eEpilobiaceae Epimediaceae Eragrostidaceae Eremolepidaceae Eremosynaceae Erica

ceae Erinaceae EriocaulaceaeEriogonaceae Eriospermaceae Erodiaceae Erycibaceae

Eryngiaceae Erysimaceae Erythroniaceae ErythropalaceaeErythrospermaceae Erythro

xylaceae Escalloniaceae Eschscholziaceae Eucomidaceae Eucommiaceae Eucryphiace

aeEuonymaceae Eupatoriaceae Euphorbiaceae Euphrasiaceae Euphroniaceae Eupo

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matiaceae Eupteleaceae EuryalaceaeEustrephaceae Euthemidaceae Eutocaceae Exb

ucklandiaceae Exocarpaceae

Fabaceae Fagaceae Falcatifoliaceae Ferulaceae Festucaceae Ficaceae Ficoidaceae Fit

zroyaceae FlacourtiaceaeFlagellariaceae Flindersiaceae Foetidiaceae Forestieraceae

Fothergillaceae Fouquieriaceae Fragariaceae FrancoaceaeFrangulaceae Frankeniacea

e Fraxinaceae Fraxinellaceae Freycinetiaceae Fritillariaceae Fuchsiaceae Fumariacea

eFunkiaceae

Gaiadendraceae Galacaceae Galanthaceae Galaxiaceae Galedupaceae Galeniaceae G

aliaceae Garciniaceae GardeniaceaeGardneriaceae Garryaceae Geissolomataceae Ge

itonoplesiaceae Gelsemiaceae Geniostomaceae GentianaceaeGeonomataceae Geosir

idaceae Geraniaceae Gerrardinaceae Gesneriaceae Gethyllidaceae Gilliesiaceae Gina

lloaceaeGinkgoaceae Gisekiaceae Gladiolaceae Glaucidiaceae Glechomaceae Glinace

ae Globulariaceae Gnaphaliaceae GnetaceaeGoetzeaceae Gomortegaceae Gomphiac

eae Gomphrenaceae Gonystylaceae Goodeniaceae Gordoniaceae GouaniaceaeGoup

iaceae Gramineae Gratiolaceae Grewiaceae Greyiaceae Grielaceae Griseliniaceae Gr

onoviaceae GrossulariaceaeGrubbiaceae Guaiacanaceae Guamatelaceae Guettardac

eae Gunneraceae Gustaviaceae Guttiferae GyrocarpaceaeGyrostemonaceae

Hachetteaceae Haemanthaceae Haemodoraceae Halesiaceae Hallieraceae Halocarpa

ceae Halophilaceae HalophytaceaeHaloragaceae Hamamelidaceae Hanguanaceae H

aptanthaceae Harmandaceae Hebenstretiaceae HectorellaceaeHederaceae Hedyosm

aceae Hedyotidaceae Hedysaraceae Heisteriaceae Heleniaceae Heliamphoraceae H

elianthaceaeHelianthemaceae Helichrysaceae Heliconiaceae Helicteraceae Heliotropi

aceae Helleboraceae Heloniadaceae HelosidaceaeHelwingiaceae Hemerocallidaceae

Hemimeridaceae Hemiphylacaceae Henriqueziaceae Henslowiaceae Hermanniaceae

Hernandiaceae Herniariaceae Herreriaceae Hesperocallidaceae Heterantheraceae He

teropyxidaceae HeterostylaceaeHibbertiaceae Hibiscaceae Hilleriaceae Himantandra

ceae Hippocastanaceae Hippocrateaceae HippomanaceaeHippophaeaceae Hippurida

ceae Hirtellaceae Holacanthaceae Homaliaceae Hopkinsiaceae Hoplestigmataceae H

ordeaceaeHornschuchiaceae Hortensiaceae Hortoniaceae Hostaceae Houstoniaceae

Huaceae Huerteaceae HugoniaceaeHumbertiaceae Humiriaceae Hyacinthaceae Hyd

atellaceae Hydnoraceae Hydrangeaceae Hydrastidaceae HydrillaceaeHydrocharitacea

e Hydrocotylaceae Hydrogetonaceae Hydroleaceae Hydropeltidaceae Hydrophylacac

eae HydrophyllaceaeHydrostachyaceae Hymenocardiaceae Hypecoaceae Hyperanth

eraceae Hypericaceae Hypopityaceae HypoxidaceaeHypseocharitaceae

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Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta

ceae ImperatoriaceaeInocarpaceae Inulaceae Ionidiaceae Iriarteaceae Iridaceae Irvi

ngiaceae Isnardiaceae Isophysidaceae IteaceaeStrasburgeriaceae Ixiaceae Ixioliriace

ae Ixonanthaceae

Jalapaceae Jamboliferaceae Japonoliriaceae Jasionaceae Jasminiaceae Johnsoniaceae

Joinvilleaceae JuglandaceaeJulianiaceae Juncaceae Juncaginaceae Juniperaceae Jus

siaeaceae Justiciaceae

Kadsuraceae Kaliphoraceae Kalmiaceae Kaniaceae Kiggelariaceae Kingdoniaceae Kin

giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae

Krameriaceae

Labiatae Lacandoniaceae Lachenaliaceae Lacistemataceae Lactoridaceae Lactucaceae

Lagerstroemiaceaee LamiaceaeLampsanaceae Lanariaceae Langsdorffiaceae Lantan

aceae Lapageriaceae Lardizabalaceae Lasiopetalaceae LauraceaeLawsoniaceae Laxm

anniaceae Lecythidaceae Ledaceae Ledocarpaceae Leeaceae Leguminosae Leitneriac

eae LemnaceaeLennoaceae Lentibulariaceae Lentiscaceae Leoniaceae Leonticaceae

Lepidobotryaceae LepidocarpaceaeLepidocaryaceae Lepidocerataceae Leptaulaceae

Leptospermaceae Lepturaceae Lepuropetalaceae LeuctenbergiaceaeLibocedraceae

Licaniaceae Ligustraceae Lilaeaceae Liliaceae Limeaceae Limnanthaceae Limnocharit

aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace

ae Linderniaceae Lindleyaceae Linnaeaceae LippayaceaeLiriaceae Liriodendraceae Li

ssocarpaceae Littorellaceae Loasaceae Lobeliaceaee Loganiaceae LomandraceaeLoni

ceraceae Lopeziaceae Lophiocarpaceae Lophiolaceae Lophiraceae Lophophytaceae L

ophopyxidaceae LoranthaceaeLotaceae Lowiaceae Lupulaceaee Luxemburgiaceae Lu

zuriagaceae Lychnidaceae Lyciaceae LyginiaceaeLygodisodeaceae Lysimachiaceae Ly

thraceae

Macarisiaceae Mackinlayaceae Madiaceae Maesaceae Magnoliaceae Malaceae Male

sherbiaceae MalortieaceaeMalpighiaceae Malvaceae Mangiaceae Manicariaceae Ma

paniaceae Marantaceae Marathraceae MarcgraviaceaeMartyniaceae Mastixiaceae

Matricariaceae Maundiaceae Mayacaceae Medeolaceae Medusagynaceae Peridiscac

eaeMelaleucaceae Melampyraceae Melanophyllaceae Melanthiaceae Melastomatac

eae Meliaceae Melianthaceae MelicaceaeMeliosmaceae Melittidaceae Melochiacea

e Memecylaceae Mendonciaceae Menispermaceae Menthaceae MenyanthaceaeMe

nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

Nypaceae Nyssaceae

Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

ceae Papilionaceae Pappophoraceae PapyraceaeParacryphiaceae Parasitaxaceae Par

ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

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hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

49

cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

50

ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 27: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

27

together may lead to increased support for clades (eg Hufford 1992 Nandi et al 1998

[but adding morphological data reduces support for a number of critical clades too]

Doyle amp Endress 2000 however in none of these papers is the use of morphology

without ambiguity) It is unfortunately clear that the use of morphology alone may not

suggest problems in the phylogenetic placement of taxa that later turn out to have been

wrongly included (eg Zhang et al 1992) Thus I have been wary of putting much weight

on clades that have only morphological support but note that I have treated molecule-

based clades with low bootstrap or jacknife support values (esp below 70) or low

posterior probabilies (below 095) likewise Although I may have been mistaken in

placing so much emphasis on molecular data in terms of providing the basic

phylogenetic framework for angiosperms morphological and molecular data are only

very rarely in irreconcilably strong conflict There are indeed a few places where the

conflict seems extreme These include the relative positions of the Monimiaceae and

Hernandiaceae (Laurales) the position of Hanguanaceae (Commelinales [as here] or

Zingiberales) and of Triplostegia (is it in Dipsacaceae or Valerianaceae - see

Dipsacales) Fossils are unlikely to affect the topologies of the trees presented here but

see below for their importance in understanding morphological evolution in general and

the evolution of angiosperms in particular Nevertheless some largely reject the idea

that trees based on molecular data alone can recover phylogenetic relationships

especially when branching points are old and prefer to used trees based on analysis of

morphological data including those taken from fossils (Hilton amp Bateman 2006 Farjon

2007)

All in all however the extent of the congruence between morphological and

molecular data is impressive and heartening and many clades can be characterised

morphologically It seemed in 1998 that there were no unambiguous morphological

synapomorphies for angiosperm orders (K Bremer 2000) and this is still true if by

unambiguous is meant non-homoplasious However many orders and other clades

have synapomorphies even if these may be indistinguishable at least at the current

level of morphological and developmental knowledge from parallel occurences

elsewhere As our morphological knowledge increases so too does the number of

apomorphies

Endress and Matthews (2006a) emphasize the importance of tendencies and

developmental constraints when thinking about characters of clades within the rosids -

many characters or character combinations occur in a rather sporadic fashion within

the clade and apparently notably less frequently outside it There are a number of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

28

examples of tendencies here and in the individual order pages such as the features

enclosed by parentheses in groups above the level of family eg see (cuticular waxes

as aggregated rodlets) for the commelinids and the discussion of the distribution of

polyandry that is flowers with numerous stamens in the asterid I + II groups etc

Tendencies may involve more than single characters When thinking of the

associations of features like integument thickness and vascularization of the integument

(Endress amp Matthews 2006a) one can imagine fairly simple developmental

preconditions being operative As Endress and Matthews (2006a) note it is difficult to

imagine vascular tissue developing in an integument only two cells thick Changes here

would be loosely correlated if morphologically linked However the strongly correlated

changes noted by Givnish et al (2005) are ecologically linked but are presumably

morphologicallydevelopmentally independent When there seem to be characters

evolving more or less together tests can be carried out to see if the changes are

concentrated on certain branches of the tree (eg Maddison 1990 Sanderson 1991

Maddison amp Maddison 2000) Such tests have rarely been carried out at higher levels in

angiosperms

One of the most striking examples of a tendency is the distribution of N-fixation

restricted as it is to a monophyletic group of four clades although it has arisen seven (or

perhaps several more) times independently within the N-fixing clade and several

members of two quite different kinds of bacteria are involved (eg Clawson et al 2004

Elliott et al 2007 Sprent amp James 2007) Other examples of work that bears on the issue

of tendencies include the findings that flowers of polysymmetrical Arabidopsis have

genes like TCP1 that are expressed asymmetrically during early development and TCP1

is a probable orthologue of the well-known CYC gene of Antirrhinum that is involved in

the development of monosymmetric slowers there (Cubas et al 2001 Costa et al 2005

etc) Parallelism might build on this underlying morphologically cryptic monosymmetry

even if details of the genetic mechanisms causing the monosymmetry evident in

particular groups may be different (Reeves amp Olmstead 2003 Cubas 2004 see above)

Similarly there have been several recent suggestions that the capability to synthesise a

particular metabolite may be switched off but not lost and so can sometimes be

reacquired (eg Wink amp Witte 1983 Wink 2003 Liscombe et al 2005 Larsson 2007

Waterman 2007) Hence perhaps the rather spotty distribution of many secondary

metabolites like ellagic acid the indole alkaloid camptothecin iridoids etc when

considered in the context of phylogenies As a non-botanical example - but a rather nice

one - Salwini-Plawen and Mayr (1961) suggested some time ago that there has been

considerable parallelism (40-65 or more independent origins) in the evolution of eyes in

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

29

metazoans However Pax 6 seems to be a master control gene that is involved in eye

formation perhaps throughout all bilateralians (eg Gehring amp Ikeo 1999 Erwin amp

Davidson 2002) which changes the nature of the problem Note however that in

general our invocation of evolutionary tendencies to explain the patterns of

distributions of characters is really an admission that we do not understand those

patterns

As we find out more about variation we will probably find fewer and fewer features

constant throughout a group Most unqualified statements of presence and absence

should properly be qualified as usually present or usually absent if one is thinking of

the characterisations on the order pages as encompassing the total variation within a

clade Thus Pistia alone among monocots as so far known has sieve tube plastids with

starch grains not protein crystals However this does not affect the fact that sieve tube

plastids with cuneate protein crystals and lacking starch grains are an apomorphy for

monocots Furthermore fossils in a number of cases suggest character combinations

unknown in extant taxa as may be seen in the discussions of Fagaceae Platanaceae

Iteaceae Calycanthaceae etc Confusing the issue there may be questions as to where

exactly on the tree a particular fossil is to be placed (eg see Nymphaeaceae

Calycanthaceae Archaefructus etc)

To summarize given our current understandings of both phylogenies and

characters evolution of some characters in which we are interested seems very labile

(see eg D Soltis et al 2005b Endress and Matthews 2006a Stevens 2006b Ekman et

al 2008 for a good recent study) and I have been cautious when talking about character

evolution Much effort must continue to be spent in summarizing characters of well-

established clades at all levels providing features by which they may be recognized and

signaling synapomorphies Remember that (1) the basic morphological anatomical and

chemical knowledge of many critical taxa is woefully incomplete (2) different

assumptions about character evolution may greatly affect the position of

synapomorphies on trees (3) in many cases relationships within and between many

groups are too uncertain at present to worry very much about synapomorphies and

(4) we must be clear about what we do and do not not know As mentioned above I

have indicated in a number of places where there are particularly important

uncertainties as to where characters should be placed on the tree Nevertheless it is a

relatively easy matter to update notes such as these and it can be a simple matter to

incorporate new data on characters that have never before been considered in the

context of a tree Much basic - and unfortunately perhaps unfashionable - work must be

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

30

carried to clarify the distribution of morphological anatomical and chemical characters

for examples of what can to be done see the work of P K Endress and collaborators

(floral morphology and development) and S R Jensen and collaborators (iridoids) But

acquiring information about nodal anatomy cell and tissue distribution nodal anatomy

and the like is not difficult at all and should be generally encouraged

But all this is merely a necesary prelude to the understanding of evolution For this

one needs to know a lot more including dating the phylogeny understanding the

function(s) of characters factoring in relevant aspects of the palaeoenvironment in

which an apomorphy first appeared etc

SUMMARY OF THE SYSTEM

Below is a formalised summary of the relationships within orders of the families of seed

plants There are a few families that are not recognised even as options in APG II and

vice versa as well as a few extra orders The families to be recognised in the most

recent edition of Mabberleys The Plant Book (Mabberley 2008) are also largely

consistent with those below But all differences are trivial and will - I hope - eventually

disappear

Square brackets - [] - enclose clades the plus sign - + - designates sister taxa a

comma - - denotes part of a polytomy and quotation marks - - denotes a

paraphyletic group

SEED PLANTS

GYMNOSPERMS

Cycadales

Cycadaceae + Zamiaceae

Ginkgoales

Ginkgoaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

31

Pinales

Pinaceae [[Araucariaceae [Podocarpaceae + Phyllocladaceae]] [Sciadopityaceae

[Taxaceae + Cupressaceae]]]

Gnetales

Ephedraceae [Gnetaceae + Welwitschiaceae]

ANGIOSPERMSFLOWERING PLANTS

Amborellales

Amborellaceae

Nymphaeales

Hydatellaceae [Cabombaceae + Nymphaeaceae]

Austrobaileyales

Austrobaileyaceae [Schisandraceae + Trimeniaceae]

Chloranthales

Chloranthaceae

MAGNOLIIDS

Magnoliales

Myristicaceae [Magnoliaceae [[Himantandraceae + Degeneriaceae] [Eupomatiaceae +

Annonaceae]]]

Laurales

Calycanthaceae [[Siparunaceae [Gomortegaceae + Atherospermataceae]] [Monimiaceae

[Hernandiaceae + Lauraceae]]]

Canellales

Canellaceae + Winteraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

32

Piperales

[Hydnoraceae + Aristolochiaceae] [Piperaceae + Saururaceae]

MONOCOTS

Acorales

Acoraceae

Alismatales

Araceae [Tofieldiaceae [[Alismataceae [Hydrocharitaceae + Butomaceae]]

[Scheuchzeriaceae [Aponogetonaceae [Juncaginaceae [Maundiaceae [[Posidoniaceae

[Ruppiaceae + Cymodoceaceae]] [Zosteraceae + Potamogetonaceae]]]]]]]

Petrosaviales

Petrosaviaceae

Dioscoreales

Nartheciaceae [[Taccaceae + Thismiaceae] [Burmanniaceae + Dioscoreaceae]]

Pandanales

Velloziaceae Triuridaceae Stemonaceae [Pandanaceae + Cyclanthaceae]

Liliales

Corsiaceae [Campynemataceae [Petermanniaceae [Colchicaceae + Alstroemeriaceae]]

Melanthiaceae [[Philesiaceae + Rhipogonaceae] [Smilacaceae + Liliaceae]]]

Asparagales

Orchidaceae [[Boryaceae [Blandfordiaceae [Lanariaceae [Asteliaceae + Hypoxidaceae]]]]

[[Ixioliriaceae + Tecophilaeaceae] [Doryanthaceae [Iridaceae [Xeronemataceae

[Xanthorrhoeaceae [Alliaceae + Asparagaceae]]]]]]]

COMMELINIDS

Unplaced

Dasypogonaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

33

Arecales

Arecaceae

Poales

[Typhaceae + Bromeliaceae] [Rapateaceae [[Thurniaceae [Juncaceae + Cyperaceae]]

[[Mayacaceae [Eriocaulaceae + Xyridaceae]] [Flagellariaceae [Anarthriaceae

[Centrolepidaceae + Restionaceae]]] [Joinvilleaceae [Ecdeiocoleaceae + Poaceae]]]]]]]

Commelinales

[Commelinaceae + Hanguanaceae] [Philydraceae [Haemodoraceae + Pontederiaceae]]

Zingiberales

Musaceae [Strelitziaceae + Lowiaceae] Heliconiaceae [[Cannaceae + Marantaceae]

[Costaceae + Zingiberaceae]]

Ceratophyllales

Ceratophyllaceae

EUDICOTS

Ranunculales

Eupteleaceae [Papaveraceae [[[Lardizabalaceae + Circaeasteraceae] [Menispermaceae

[Berberidaceae + Ranunculaceae]]]]

Sabiales

Sabiaceae

Proteales

Nelumbonaceae [Platanaceae + Proteaceae]

Trochodendrales

Trochodendraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

34

Buxales

Haptanthaceae [Buxaceae + Didymelaceae]

CORE EUDICOTS

Gunnerales

Gunneraceae + Myrothamnaceae

Dilleniales

Dilleniaceae

Saxifragales

Peridiscaceae [[Paeoniaceae [Altingiaceae [Hamamelidaceae [Cercidiphyllaceae +

Daphniphyllaceae]]]] [[Crassulaceae [Aphanopetalaceae [Tetracarpaeaceae

[Penthoraceae + Haloragaceae]]]] [Iteaceae [Grossulariaceae + Saxifragaceae]]]]

Cynomoriaceae unplaced

Vitales

Vitaceae

ROSIDS

FABIDROSID I

Zygophyllales

Krameriaceae + Zygophyllaceae

Celastrales

Lepidobotryaceae + Celastraceae

Oxalidales

Huaceae [[Connaraceae + Oxalidaceae] [Cunoniaceae [Elaeocarpaceae [Brunelliaceae +

Cephalotaceae]]]]

Malpighiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

35

[Achariaceae [Goupiaceae [Violaceae + Passifloraceae] [Lacistemataceae Salicaceae]]

[Lophopyxidaceae Putranjivaceae] [Ctenolophonaceae [Erythroxylaceae +

Rhizophoraceae]] Linaceae Ixonanthaceae Humiriaceae Irvingiaceae

Centroplacaceae Pandaceae [[Bonnetiaceae + Clusiaceae] [Calophyllaceae

[Hypericaceae + Podostemaceae]]] [Malpighiaceae + Elatinaceae] Ochnaceae

[Peraceae [Rafflesiaceae + Euphorbiaceae]] [Phyllanthaceae + Picrodendraceae]

[Balanopaceae [[Trigoniaceae + Dichapetalaceae] [Chrysobalanaceae +

Euphroniaceae]]] Caryocaraceae

N-FIXING CLADE

Fabales

Quillajaceae [Fabaceae [Polygalaceae + Surianaceae]]

Rosales

Rosaceae [[Barbeyaceae [Elaeagnaceae Dirachmaceae Rhamnaceae]] [Ulmaceae

[Cannabaceae [Moraceae + Urticaceae]]]]

Cucurbitales

Anisophylleaceae [[Corynocarpaceae + Coriariaceae] [Cucurbitaceae [Tetramelaceae

[Datiscaceae + Begoniaceae]]]] Apodanthaceae

Fagales

Nothofagaceae [Fagaceae [[Myricaceae + Juglandaceae] [Casuarinaceae

[Ticodendraceae + Betulaceae]]]]

MALVIDROSID II

pgtGeraniales

Geraniaceae [[Melianthaceae + Francoaceae] [Vivianaceae + Ledocarpaceae]]

Myrtales

Combretaceae [[Onagraceae + Lythraceae] [[Vochysiaceae + Myrtaceae]

[Melastomataceae [Crypteroniaceae [Alzateaceae + Penaeaceae]]]]]

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

36

Crossosomatales

[Staphyleaceae [Guamatelaceae [Crossosomataceae + Stachyuraceae]]] [Aphloiaceae

[Geissolomataceae + Strasburgeriaceae]]

Picramniales

Picramniaceae

Sapindales

Biebersteiniaceae Nitrariaceae [Kirkiaceae [Anacardiaceae + Burseraceae]]

[Sapindaceae [Simaroubaceae Rutaceae Meliaceae]]

Huerteales

Gerradinaceae [Dipentodontaceae + Tapisciaceae]

Malvales

Neuradaceae [Thymelaeaceae [Sphaerosepalaceae Bixaceae [Cistaceae

[Sarcolaenaceae + Dipterocarpaceae]] [Cytinaceae + Muntingiaceae] Malvaceae]]

Brassicales

[Akaniaceae + Tropaeolaceae] [[Moringaceae + Caricaceae] [Setchellanthaceae

[Limnanthaceae [[Koeberliniaceae [Bataceae + Salvadoraceae]] [Emblingiaceae

[Pentadiplandraceae [Gyrostemonaceae Resedaceae Stixis and relatives] Tovariaceae

[Capparaceae [Cleomaceae + Brassicaceae]]]]]]]]

Berberidopsidales

Aextoxicaceae + Berberidopsidaceae

Santalales

Erythropalaceae Olacaceae [[Loranthaceae [Misodendraceae + Schoepfiaceae]]

[Opiliaceae + Santalaceae]] Balanophoraceae unplaced

Caryophyllales

[[Droseraceae [Nepenthaceae [Drosophyllaceae [Ancistrocladaceae +

Dioncophylleaceae]]]]] [[Frankeniaceae + Tamaricaceae] [Polygonaceae +

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

37

Plumbaginaceae]]] [Rhabdodendraceae [Simmondsiaceae [[Asteropeiaceae +

Physenaceae] [[Caryophyllaceae [Achatocarpaceae + Amaranthaceae]]

[Stegnospermataceae [Limeaceae [[Lophocarpaceae [Barbeuiaceae [Aizoaceae

[Sarcobataceae Phytolaccaceae Nyctaginaceae]]]] [Molluginaceae [Halophytaceae

Montiaceae Basellaceae Didiereaceae [Talinaceae [Portulacaceae [Anacampseros etc +

Cactaceae]]]]]]]]]]]]]

ASTERIDS

Cornales

[Cornaceae + Nyssaceae] [Hydrangeaceae + Loasaceae] Hydrostachyaceae [Curtisiaceae

+ Grubbiaceae]

Ericales

[Balsaminaceae [Marcgraviaceae + Tetrameristaceae]] [[Polemoniaceae +

Fouquieraceae] Lecythidaceae [[Sladeniaceae + Pentaphylacaceae] [Sapotaceae

[Ebenaceae [Maesaceae [Theophrastaceae [Myrsinaceae + Primulaceae]]]]]

[Mitrastemonaceae Theaceae [Symplocaceae [Styracaceae + Diapensiaceae]]

[[Sarraceniaceae [Roridulaceae + Actinidiaceae] [Clethraceae [Cyrillaceae +

Ericaceae]]]]]]

LAMIIDASTERID I

Unplaced

Metteniusaceae Oncothecaceae Icacinaceae and some unplaced ex-Icacinaceae

Garryales

Garryaceae + Eucommiaceae

Unplaced

Boraginaceae Vahliaceae

Gentianales

Rubiaceae [Gentianaceae [Loganiaceae [Gelsemiaceae + Apocynaceae]]]

Lamiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

38

Plocospermataceae [Oleaceae [Tetrachondraceae [Calceolariaceae

[Gesneriaceae Peltanthera [Pedaliaceae Carlemanniaceae Martyniaceae Stilbaceae

Plantaginaceae Linderniaceae Bignoniaceae Verbenaceae Lamiaceae Paulowniaceae

Schlegeliaceae Thomandersiaceae Phrymaceae Rehmannia Acanthaceae

Scrophulariaceae Orobanchaceae Byblidaceae Lentibulariaceae]]]]]

Solanales

[Montiniaceae [Sphenocleaceae + Hydroleaceae]] [Convolvulaceae + Solanaceae]

CAMPANULIDASTERID II

Aquifoliales

[[Cardiopteridaceae + Stemonuraceae] [Aquifoliaceae [Helwingiaceae +

Phyllonomaceae]]

Asterales

[Rousseaceae + Campanulaceae] Pentaphragmataceae [[Alseuosmiaceae [Phellinaceae

+ Argophyllaceae]] Stylidiaceae [Menyanthaceae [Goodeniaceae [Calyceraceae +

Asteraceae]]]]

Escalloniales

Escalloniaceae

Bruniales

[Bruniaceae + Columelliaceae]

Apiales

Pennantiaceae [Torricelliaceae [Griseliniaceae [Pittosporaceae [Araliaceae

[Myodocarpaceae + Apiaceae]]]]]

Paracryphiales

Paracryphiaceae

Dipsacales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

39

Adoxaceae [Diervillaceae [Caprifoliaceae [Linnaeaceae [Morinaceae [Dipsacaceae +

Valerianaceae]]]]]

LINKS TO ORDERS AND FAMILIES

[Back to Top]

MAIN GROUPINGS

Asterids commelinids core eudicots asterid 1 asterid

2 eudicots gymnosperms Magnoliophyta monocots N-fixing claderosids seed plants

ALPHABETICAL LISTING OF ALL ORDINAL NAMES OF SEED PLANTS WITH LINKS

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Abietales Acanthales Acerales Acorales Actinidiales Actinostrobales Adoxales Aescul

ales Agavales Aizoales AkanialesAlismatales Alliales Alseuosmiales Alstroemeriales

Altingiales Amaranthales Amaryllidales Amborellales AmbrosialesAmmiales Amomal

es Ancistrocladales Anisophylleales Annonales Anthobolales Apiales Apocynales Apo

nogetonalesAquifoliales Arales Araliales Aralidiales Araucariales Arecales Aristolochi

ales Athrotaxidales Asarales AsclepiadalesAsparagales Asphodelales Asparagales Ast

eliales Atriplicales Aucubales Austrobaileyales Avenales

Balanitales Balanopales Balanophorales Balsaminales Barbeyales Barclayales Batales

Begoniales BerberidalesBerberidopsidales Betulales Biebersteiniales Bignoniales Bix

ales Boraginales Brassicales Brexiales Bromeliales BrunialesBrunoniales Burmannial

es Burserales Butomales Buxales Byblidales

Cactales Callitrichales Calycanthales Calycerales Campanulales Campynematales Can

ellales Cannales CapparalesCaprifoliales Cardiopteridales Carduales Caricales Carle

manniales Caryophyllales Cassiales Casuarinales CelastralesCentrolepidales Cephalot

ales Cephalotaxales Ceratophyllales Cercidiphyllales Chenopodiales Chironiales Chlor

anthalesChrysobalanales Cinchonales Circaeasterales Cistales Citrales Cocosales Col

chicales Columelliales CombretalesCommelinales Connarales Convolvulales Coriarial

es Cornales Corylales Corynocarpales Crassulales CrossosomatalesCucurbitales Cunn

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

40

inghamiales Cunoniales Cupressales Cycadales Cyclanthales Cymodoceales Cynarales

CynomorialesCyperales Cyrillales Cytinales

Daphnales Daphniphyllales Dasypogonales Datiscales Degeneriales Desfontainiales D

iapensiales Didymelales DillenialesDioales Dioncophyllales Dioscoreales Diospyrales

Dipentodontales Dipsacales Droserales

Ebenales Echiales Elaeagnales Elaeocarpales Elatinales Elodeales Emmotales Empetr

ales Ephedrales Ericales EriocaulalesErythropalaless Escalloniales Eucommiales Euph

orbiales Eupomatiales Eupteleales Euryalaless

Fabales Fagales Falcatifoliales Ficales Flacourtiales Flagellariales Fouquieriales Franc

oales Frangulales

Galiales Garryales Geissolomatales Gentianales Geraniales Gesneriales Ginkgoales G

laucidiales Globulariales GnetalesGoodeniales Greyiales Griseliniales Grossulariales

Grubbiales Gunnerales Gyrocarpales Gyrostemonales

Haemodorales Haloragales Hamamelidales Hanguanales Heisteriales Helleborales He

lwingiales Himantandrales HippuridalesHomaliales Hortensiales Huales Huerteales

Hydatellales Hydnorales Hydrangeales Hydrastidales HydrocharitalesHydropeltidales

Hydrostachyales Hypericales Hypoxidales

Icacinales Illiciales Iridales Irvingales Iteales Ixiales

Jasminales Juglandales Julianiales Juncaginales Juncales

Lacistematales Lactoridales Lamiales Lardizabalales Laurales Lecythidales Ledocarpal

es Leitneriales LentibularialesLigustrales Liliales Limnanthales Linales Liriales Loasal

es Lobeliales Loganiales Lonicerales Loranthales LowialesLythrales

Magnoliales Malpighiales Malvales Marathrales Marcgraviales Mayacales Medusagy

nales Medusandrales MelanthialesMelastomatales Meliales Melianthales Meliosmal

es Menispermales Menyanthales Metteniusales MitrastemonalesMiyoshiales Moni

miales Moringales Musales Myricales Myristicales Myrothamnales Myrsinales Myrta

les

Najadales Nandinales Narcissales Nartheciales Nelumbonales Nepenthales Neuradal

es Nitrariales Nolanales NothofagalesNyctaginales Nymphaeales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

41

Ochnales Oenotherales Olacales Oleales Onagrales Oncothecales Opuntiales Orchid

ales Oxalidales

Saxifragales Pandales Pandanales Papaverales Paracryphiales Parasitaxales Paridales

Parnassiales PassifloralesPenaeales Pennantiales Pentaphragmatales Peridiscales P

etiveriales Petrosaviales Phellinales Philydrales PhyllanthalesPhysenales Phytolaccal

es Picramniales Pinales Pinguiculales Piperales Pittosporales Plantaginales Platanale

sPlumbaginales Poales Podocarpales Podophyllales Podostemales Polemoniales Pol

ygalales Polygonales PontederialesPortulacales Posidoniales Potamogetonales Primu

lales Proteales

Quercales Quillajales Quintiniales

Rafflesiales Ranunculales Rapateales Resedales Restionales Rhabdodendrales Rhamn

ales Rhinanthales RhizophoralesRhodorales Rhoipteleales Roridulales Rosales Rouss

eales Rubiales Ruppiales Rutales

Sabiales Salicales Salvadorales Samolales Samydales Sanguisorbales Santalales Sapin

dales Sapotales SarracenialesSaxegotheales Saxifragales Scheuchzeriales Sciadopitya

les Scleranthales Scrophulariales Scyphostegiales SedalesSilenales Simmondsiales S

milacales Solanales Sphenocleales Sphenostemonales Stangeriales Stellariales Stemo

nalesStilbales Stylidiales Styracales Surianales

Taccales Tamales Tamaricales Taxales Taxodiales Tecophilaeales Terebinthales Tern

stroemiales Theales TheligonalesThymelaeales Tiliales Tofieldiales Torricelliales Tov

ariales Tribelales Trilliales Trimeniales Triuridales TrochodendralesTropaeolales Turn

erales Typhales

Ulmales Urticales

Vacciniales Vahliales Vallisneriales Velloziales Veratrales Verbenales Viburnales Vinc

ales Violales Viscales VitalesVochysiales

Welwitschiales Winterales

Xanthorrhoeales Ximeniales Xyridales

Zamiales Zingiberales Zosterales Zygophyllales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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ALPHABETICAL LISTING OF ALL FAMILY NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | K | L | M | N | O | P | Q | R | S | T | U | V | W | X | Y

|Z

Back to Home

In some cases clicking on a family synonym will send you directly to the subfamily in

which the genus of the basionym is to be found

Abietaceae Abolbodaceae Abrophyllaceae Acalyphaceae Acanthaceae [Acanthochla

mydaceae] Acarnaceaee AceraceaeAchariaceae Achatocarpaceae Achradaceae Achy

ranthaceae Acmopylaceae Acoraceae Acristaceae ActaeaceaeActinidiaceae Actinota

ceae Amentotaxaceae Adenogrammaceae Adoxaceae Aegialitidaceae Aegicerataceae

Aegilopaceae Aeginetiaceaee Aegiphilaceae Aesculaceae Aextoxicaceae Agapanthac

eae Agavaceae AgdestidaceaeAgialidaceae Agrimoniaceae Agrostidaceae Ailanthace

ae Aitoniaceae Aizoaceae Akaniaceae Alangiaceae AlchemillaceaeAldrovandaceae A

lismataceae Alliaceae Allioniaceae Allophylaceae Aloaceae Alopecuraceae Alpiniacea

e AlseuosmiaceaeAlsinaceae Alsodeiaceae Alstroemeriaceae Altingiaceae Alzateacea

e Amaranthaceae Amaryllidaceae Amborellaceae Ambrosiaceae Amentotaxaceae A

mmanniaceae Ammiaceae Amomaceae Ampelidopsaceae Amygdalaceae Amyridacea

eAnacardiaceae Anagallidaceae Anarthriaceae Anchusaceae Ancistrocladaceae Andr

omedaceae AndropogonaceaeAndrostachyaceae Androsynaceae Anemarrhenaceae

Anemonaceae Angelicaceae Anisophylleaceae AnnonaceaeAnomochloaceae Anopter

aceae Anrederaceae Anthemidaceae Anthericaceae Anthobolaceae Antidesmataceae

Antirrhinaceae Antoniaceae Aparinaceae Aphanopetalaceae Aphloiaceae Aphyllanth

aceae Apiaceae ApocynaceaeApodanthaceae Aponogetonaceae Aporusaceae Apose

ridaceae Apostasiaceae Aptandraceae AquifoliaceaeAquilariaceae Aquilegiaceae A

raceae Aragoaceae Araliaceae Aralidiaceaee Araucariaceae Arbutaceae Arceuthidace

aeArceuthobiaceae Arctostaphylaceae Arctotidaceae Ardisiaceae Arecaceae Argoph

yllaceae Arisaraceae AristolochiaceaeAristoteliaceae Arjonaceae Armeriaceae Artem

isiaceae Arthrotaxidaceae Artocarpaceae Arundinaceae ArundinellaceaeAsaraceae A

sclepiadaceae Ascyraceaee Asiraceae Asparagaceae Asperulaceae Asphodelaceae As

pidistraceaeeAsteliaceae Asteraceae Asteranthaceae Asteropeiaceae Astragalaceae

Astrocarpaceae AthanasiaceaeAtherospermataceae Atriplicaceae Atropaceae Aucuba

ceae Aurantiaceae Austrobaileyaceae Austrotaxaceae AvenaceaeAverrhoaceae Avet

raceae Avicenniaceae Azaleaceae Azimaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

43

Balanitaceae Balanopaceae Balanophoraceae Balsameaceae Balsaminaceae Bambusa

ceae Barbaceniaceae BarbeuiaceaeBarbeyaceae Barclayaceae Barringtoniaceae Base

llaceae Bataceae Baueraceae Bauhiniaceae Baxteriaceae BegoniaceaeBehniaceae B

elangeraceae Belloniaceae Bembiciaceae Berberidaceae Berberidopsidaceae Berryac

eaee BersamaceaeBertyaceae Berzeliaceae Besleriaceae Betaceae Betulaceae Biebe

rsteiniaceae Bifariaceae Bignoniaceae BischofiaceaeBixaceae Blakeaceae Blandfordia

ceae Blattiaceae Blepharocaryaceae Blitaceae Blyxaceae Boerlagellaceae Bolivaracea

eBombacaceae Bonnetiaceae Bontiaceae Boopidaceae Boraginaceae Borassaceae B

oroniaceae BoryaceaeBotryodendraceae Bougainvilleaceae Boweniaceae Brassicacea

e Brachycaulaceae Bretschneideraceae BrexiaceaeBromeliaceae Brownlowiaceae Br

unelliaceae Bruniaceae Brunoniaceae Brunsvigiaceae Bryoniaceae BuchneraceaeBuci

daceae Buddlejaceae Buglossaceae Bulbocodiaceae Bumeliaceae Bupleuraceae Burc

hardiaceae BurmanniaceaeBurseraceae Butneriaceae Butomaceae Buxaceae Byblida

ceae Byttneriaceae

Cabombaceae Cacaoaceae Cactaceae Caesalpiniaceae Caladiaceae Calamaceae Calce

olariaceae CalectasiaceaeCalendulaceae Callaceae Callicomaceae Calligonaceae Calli

traceae Callitrichaceae Calochortaceae CalophyllaceaeCalthaceae Calycanthaceae Ca

lyceraceae Cambogiaceae Camelliaceae Campanulaceae CampynemataceaeCanacom

yricaceae Canellaceae Cannabaceae Cannaceae Canopodaceae Canotiaceae Cansjera

ceae CapparaceaeCaprariaceae Caprifoliaceae Cardiopteridaceae Carduaceae Carica

ceae Carissaceae Carlemanniaceae CarpinaceaeCarpodetaceae Cartonemataceae Ca

ryocaraceae Caryophyllaceae Casearieaceae Cassiaceae CassipoureaceaeCassythacea

e Castaneaceae Castelaceae Casuarinaceae Catesbaeaceae Cathedraceae Cecropiace

ae CedraceaeCedrelaceae Celastraceae Celosiaceae Celtidaceae Centaureaceae Cen

trolepidaceae Centroplacaceae CepaceaeCephalanthaceae Cephalotaceae Cephalota

xaceae Cerastiaceae Ceratoniaceae Ceratophyllaceae CerberaceaeCercidiphyllaceae

Cercocarpaceae Cercodiaceae Cereaceae Cerinthaceae Ceroxylaceae Cestraceae Cev

alliaceaeChailletiaceae Chamaedoraceae Chamaemoraceae Chamelauciaceae Chaun

ochitonaceae Chelidoniaceae ChelonaceaeChenopodiaceae Chimonanthaceae Chingi

thamnaceae Chionographidaceae Chiranthodendraceae ChironiaceaeChlaenaceae Ch

loanthaceae Chloranthaceae Chloridaceae Chlorogalaceae Chrysobalanaceae Cicerac

eae CichoriaceaeCimicifugaceae Cinchoniaceae Circaeaceae Circaeasteraceae Cissac

eae Cistaceae Citraceae Clematidaceae CleomaceaeClethraceae Clusiaceae Cneorac

eae Cnestidaceae Cnicaceae Cobaeaceae Cochlospermaceae Cocosaceae Coffeaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Colchicaceae Coleogynaceae Columelliaceae Combretaceae Commelinaceae Comocl

adiaceae Compositae CompsoaceaeConantheraceae Connaraceae Conostylidaceae C

onvallariaceae Convolvulaceae Cordiaceae CoreopsidaceaeCoriandraceae Coriariacea

e Coridaceae Corispermaceae Cornaceae Corokiaceae Coronillaceae Correaceae Cor

rigiolaceaeCorsiaceae Corydalaceae Corylaceae Corynocarpaceae Coryphaceae Cost

aceae Cotyledonaceae CoulaceaeCoutariaceae Coutoubiaceae Crassulaceae Crescen

tiaceae Cressaceae Crinaceae Crocaceae CroomiaceaeCrossosomataceae Crotonacea

e Cruciferae Cryptaceae Crypteroniaceae Cryptocorynaceae CryptomeriaceaeCtenol

ophonaceae Cucurbitaceae Cunninghamiaceae Cunoniaceae Cupressaceae Curcumac

eae Curtisiaceae CuscutaceaeCyananthaceae Cyanastraceae Cynanchaceae Cyanella

ceae Cycadaceae Cyclanthaceae CyclantheraceaeCyclocheilaceae Cydoniaceae Cymo

doceaceae Cynaraceae Cynocrambaceae Cynomoriaceae Cyperaceae CyphiaceaeCyp

hocarpaceae Cypripediaceae Cyrillaceae Cyrtandraceae Cyrtanthaceae Cytinaceae

Dacrycarpaceae Dactylanthaceae Damasoniaceae Daphnaceae Daphniphyllaceae Das

ypogonaceae DatiscaceaeDaturaceae Daucaceae Davidiaceae Davidsoniaceae Decais

neaceae Deeringiaceae Degeneriaceae DelphiniaceaeDendrophthoaceae Columelliac

eae Detariaceae Dialypetalanthaceae Dianellaceae Dianthaceae DiapensiaceaeDicha

petalaceae Dichondraceae Diclidantheraceae Dicrastylidiaceae Dictamnaceae Didiere

aceae DidymelaceaeDidymocarpaceae Diegodendraceae Diervillaceae Digitalidaceae

e Dilatridaceae Dilleniaceae Dioaceae DionaeaceaeDioncophyllaceae Dioscoreaceae

Diosmaceae Diospyraceae Dipentodontaceae Diphylleiaceae Diplolaenaceae Dipsaca

ceaeDipterocarpaceae Dirachmaceae Disanthaceae Diselmaceae Dodonaeaceae Do

mbeyaceae Donatiaceae DorsteniaceaeDortmannaceae Doryanthaceae Drabaceae D

racaenaceae Dracontiaceae Drimyidaceae Droseraceae DrosophyllaceaeDryadaceae

Duabangaceae Duckeodendraceae Dulongiaceae Durantaceae Durionaceae Dysphani

aceae

Ebenaceae Eccremidaceae Ecdeiocoleaceae Echiaceae Echinopaceae Ehretiaceae Ela

eagnaceae ElaeocarpaceaeElegiaceae Elatinaceae Ellisiaceae Ellisiophyllaceae Elodea

ceae Elytranthaceae Embeliaceae Emblingiaceae EmmotaceaeEmpetraceae Encepha

lartaceae Engelhardtiaceae Enhalaceae Epacridaceae Ephedraceae Ephemeranthacea

eEpilobiaceae Epimediaceae Eragrostidaceae Eremolepidaceae Eremosynaceae Erica

ceae Erinaceae EriocaulaceaeEriogonaceae Eriospermaceae Erodiaceae Erycibaceae

Eryngiaceae Erysimaceae Erythroniaceae ErythropalaceaeErythrospermaceae Erythro

xylaceae Escalloniaceae Eschscholziaceae Eucomidaceae Eucommiaceae Eucryphiace

aeEuonymaceae Eupatoriaceae Euphorbiaceae Euphrasiaceae Euphroniaceae Eupo

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

45

matiaceae Eupteleaceae EuryalaceaeEustrephaceae Euthemidaceae Eutocaceae Exb

ucklandiaceae Exocarpaceae

Fabaceae Fagaceae Falcatifoliaceae Ferulaceae Festucaceae Ficaceae Ficoidaceae Fit

zroyaceae FlacourtiaceaeFlagellariaceae Flindersiaceae Foetidiaceae Forestieraceae

Fothergillaceae Fouquieriaceae Fragariaceae FrancoaceaeFrangulaceae Frankeniacea

e Fraxinaceae Fraxinellaceae Freycinetiaceae Fritillariaceae Fuchsiaceae Fumariacea

eFunkiaceae

Gaiadendraceae Galacaceae Galanthaceae Galaxiaceae Galedupaceae Galeniaceae G

aliaceae Garciniaceae GardeniaceaeGardneriaceae Garryaceae Geissolomataceae Ge

itonoplesiaceae Gelsemiaceae Geniostomaceae GentianaceaeGeonomataceae Geosir

idaceae Geraniaceae Gerrardinaceae Gesneriaceae Gethyllidaceae Gilliesiaceae Gina

lloaceaeGinkgoaceae Gisekiaceae Gladiolaceae Glaucidiaceae Glechomaceae Glinace

ae Globulariaceae Gnaphaliaceae GnetaceaeGoetzeaceae Gomortegaceae Gomphiac

eae Gomphrenaceae Gonystylaceae Goodeniaceae Gordoniaceae GouaniaceaeGoup

iaceae Gramineae Gratiolaceae Grewiaceae Greyiaceae Grielaceae Griseliniaceae Gr

onoviaceae GrossulariaceaeGrubbiaceae Guaiacanaceae Guamatelaceae Guettardac

eae Gunneraceae Gustaviaceae Guttiferae GyrocarpaceaeGyrostemonaceae

Hachetteaceae Haemanthaceae Haemodoraceae Halesiaceae Hallieraceae Halocarpa

ceae Halophilaceae HalophytaceaeHaloragaceae Hamamelidaceae Hanguanaceae H

aptanthaceae Harmandaceae Hebenstretiaceae HectorellaceaeHederaceae Hedyosm

aceae Hedyotidaceae Hedysaraceae Heisteriaceae Heleniaceae Heliamphoraceae H

elianthaceaeHelianthemaceae Helichrysaceae Heliconiaceae Helicteraceae Heliotropi

aceae Helleboraceae Heloniadaceae HelosidaceaeHelwingiaceae Hemerocallidaceae

Hemimeridaceae Hemiphylacaceae Henriqueziaceae Henslowiaceae Hermanniaceae

Hernandiaceae Herniariaceae Herreriaceae Hesperocallidaceae Heterantheraceae He

teropyxidaceae HeterostylaceaeHibbertiaceae Hibiscaceae Hilleriaceae Himantandra

ceae Hippocastanaceae Hippocrateaceae HippomanaceaeHippophaeaceae Hippurida

ceae Hirtellaceae Holacanthaceae Homaliaceae Hopkinsiaceae Hoplestigmataceae H

ordeaceaeHornschuchiaceae Hortensiaceae Hortoniaceae Hostaceae Houstoniaceae

Huaceae Huerteaceae HugoniaceaeHumbertiaceae Humiriaceae Hyacinthaceae Hyd

atellaceae Hydnoraceae Hydrangeaceae Hydrastidaceae HydrillaceaeHydrocharitacea

e Hydrocotylaceae Hydrogetonaceae Hydroleaceae Hydropeltidaceae Hydrophylacac

eae HydrophyllaceaeHydrostachyaceae Hymenocardiaceae Hypecoaceae Hyperanth

eraceae Hypericaceae Hypopityaceae HypoxidaceaeHypseocharitaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta

ceae ImperatoriaceaeInocarpaceae Inulaceae Ionidiaceae Iriarteaceae Iridaceae Irvi

ngiaceae Isnardiaceae Isophysidaceae IteaceaeStrasburgeriaceae Ixiaceae Ixioliriace

ae Ixonanthaceae

Jalapaceae Jamboliferaceae Japonoliriaceae Jasionaceae Jasminiaceae Johnsoniaceae

Joinvilleaceae JuglandaceaeJulianiaceae Juncaceae Juncaginaceae Juniperaceae Jus

siaeaceae Justiciaceae

Kadsuraceae Kaliphoraceae Kalmiaceae Kaniaceae Kiggelariaceae Kingdoniaceae Kin

giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae

Krameriaceae

Labiatae Lacandoniaceae Lachenaliaceae Lacistemataceae Lactoridaceae Lactucaceae

Lagerstroemiaceaee LamiaceaeLampsanaceae Lanariaceae Langsdorffiaceae Lantan

aceae Lapageriaceae Lardizabalaceae Lasiopetalaceae LauraceaeLawsoniaceae Laxm

anniaceae Lecythidaceae Ledaceae Ledocarpaceae Leeaceae Leguminosae Leitneriac

eae LemnaceaeLennoaceae Lentibulariaceae Lentiscaceae Leoniaceae Leonticaceae

Lepidobotryaceae LepidocarpaceaeLepidocaryaceae Lepidocerataceae Leptaulaceae

Leptospermaceae Lepturaceae Lepuropetalaceae LeuctenbergiaceaeLibocedraceae

Licaniaceae Ligustraceae Lilaeaceae Liliaceae Limeaceae Limnanthaceae Limnocharit

aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace

ae Linderniaceae Lindleyaceae Linnaeaceae LippayaceaeLiriaceae Liriodendraceae Li

ssocarpaceae Littorellaceae Loasaceae Lobeliaceaee Loganiaceae LomandraceaeLoni

ceraceae Lopeziaceae Lophiocarpaceae Lophiolaceae Lophiraceae Lophophytaceae L

ophopyxidaceae LoranthaceaeLotaceae Lowiaceae Lupulaceaee Luxemburgiaceae Lu

zuriagaceae Lychnidaceae Lyciaceae LyginiaceaeLygodisodeaceae Lysimachiaceae Ly

thraceae

Macarisiaceae Mackinlayaceae Madiaceae Maesaceae Magnoliaceae Malaceae Male

sherbiaceae MalortieaceaeMalpighiaceae Malvaceae Mangiaceae Manicariaceae Ma

paniaceae Marantaceae Marathraceae MarcgraviaceaeMartyniaceae Mastixiaceae

Matricariaceae Maundiaceae Mayacaceae Medeolaceae Medusagynaceae Peridiscac

eaeMelaleucaceae Melampyraceae Melanophyllaceae Melanthiaceae Melastomatac

eae Meliaceae Melianthaceae MelicaceaeMeliosmaceae Melittidaceae Melochiacea

e Memecylaceae Mendonciaceae Menispermaceae Menthaceae MenyanthaceaeMe

nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

47

Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

Nypaceae Nyssaceae

Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

ceae Papilionaceae Pappophoraceae PapyraceaeParacryphiaceae Parasitaxaceae Par

ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

48

hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

49

cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

50

ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 28: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

28

examples of tendencies here and in the individual order pages such as the features

enclosed by parentheses in groups above the level of family eg see (cuticular waxes

as aggregated rodlets) for the commelinids and the discussion of the distribution of

polyandry that is flowers with numerous stamens in the asterid I + II groups etc

Tendencies may involve more than single characters When thinking of the

associations of features like integument thickness and vascularization of the integument

(Endress amp Matthews 2006a) one can imagine fairly simple developmental

preconditions being operative As Endress and Matthews (2006a) note it is difficult to

imagine vascular tissue developing in an integument only two cells thick Changes here

would be loosely correlated if morphologically linked However the strongly correlated

changes noted by Givnish et al (2005) are ecologically linked but are presumably

morphologicallydevelopmentally independent When there seem to be characters

evolving more or less together tests can be carried out to see if the changes are

concentrated on certain branches of the tree (eg Maddison 1990 Sanderson 1991

Maddison amp Maddison 2000) Such tests have rarely been carried out at higher levels in

angiosperms

One of the most striking examples of a tendency is the distribution of N-fixation

restricted as it is to a monophyletic group of four clades although it has arisen seven (or

perhaps several more) times independently within the N-fixing clade and several

members of two quite different kinds of bacteria are involved (eg Clawson et al 2004

Elliott et al 2007 Sprent amp James 2007) Other examples of work that bears on the issue

of tendencies include the findings that flowers of polysymmetrical Arabidopsis have

genes like TCP1 that are expressed asymmetrically during early development and TCP1

is a probable orthologue of the well-known CYC gene of Antirrhinum that is involved in

the development of monosymmetric slowers there (Cubas et al 2001 Costa et al 2005

etc) Parallelism might build on this underlying morphologically cryptic monosymmetry

even if details of the genetic mechanisms causing the monosymmetry evident in

particular groups may be different (Reeves amp Olmstead 2003 Cubas 2004 see above)

Similarly there have been several recent suggestions that the capability to synthesise a

particular metabolite may be switched off but not lost and so can sometimes be

reacquired (eg Wink amp Witte 1983 Wink 2003 Liscombe et al 2005 Larsson 2007

Waterman 2007) Hence perhaps the rather spotty distribution of many secondary

metabolites like ellagic acid the indole alkaloid camptothecin iridoids etc when

considered in the context of phylogenies As a non-botanical example - but a rather nice

one - Salwini-Plawen and Mayr (1961) suggested some time ago that there has been

considerable parallelism (40-65 or more independent origins) in the evolution of eyes in

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

29

metazoans However Pax 6 seems to be a master control gene that is involved in eye

formation perhaps throughout all bilateralians (eg Gehring amp Ikeo 1999 Erwin amp

Davidson 2002) which changes the nature of the problem Note however that in

general our invocation of evolutionary tendencies to explain the patterns of

distributions of characters is really an admission that we do not understand those

patterns

As we find out more about variation we will probably find fewer and fewer features

constant throughout a group Most unqualified statements of presence and absence

should properly be qualified as usually present or usually absent if one is thinking of

the characterisations on the order pages as encompassing the total variation within a

clade Thus Pistia alone among monocots as so far known has sieve tube plastids with

starch grains not protein crystals However this does not affect the fact that sieve tube

plastids with cuneate protein crystals and lacking starch grains are an apomorphy for

monocots Furthermore fossils in a number of cases suggest character combinations

unknown in extant taxa as may be seen in the discussions of Fagaceae Platanaceae

Iteaceae Calycanthaceae etc Confusing the issue there may be questions as to where

exactly on the tree a particular fossil is to be placed (eg see Nymphaeaceae

Calycanthaceae Archaefructus etc)

To summarize given our current understandings of both phylogenies and

characters evolution of some characters in which we are interested seems very labile

(see eg D Soltis et al 2005b Endress and Matthews 2006a Stevens 2006b Ekman et

al 2008 for a good recent study) and I have been cautious when talking about character

evolution Much effort must continue to be spent in summarizing characters of well-

established clades at all levels providing features by which they may be recognized and

signaling synapomorphies Remember that (1) the basic morphological anatomical and

chemical knowledge of many critical taxa is woefully incomplete (2) different

assumptions about character evolution may greatly affect the position of

synapomorphies on trees (3) in many cases relationships within and between many

groups are too uncertain at present to worry very much about synapomorphies and

(4) we must be clear about what we do and do not not know As mentioned above I

have indicated in a number of places where there are particularly important

uncertainties as to where characters should be placed on the tree Nevertheless it is a

relatively easy matter to update notes such as these and it can be a simple matter to

incorporate new data on characters that have never before been considered in the

context of a tree Much basic - and unfortunately perhaps unfashionable - work must be

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

30

carried to clarify the distribution of morphological anatomical and chemical characters

for examples of what can to be done see the work of P K Endress and collaborators

(floral morphology and development) and S R Jensen and collaborators (iridoids) But

acquiring information about nodal anatomy cell and tissue distribution nodal anatomy

and the like is not difficult at all and should be generally encouraged

But all this is merely a necesary prelude to the understanding of evolution For this

one needs to know a lot more including dating the phylogeny understanding the

function(s) of characters factoring in relevant aspects of the palaeoenvironment in

which an apomorphy first appeared etc

SUMMARY OF THE SYSTEM

Below is a formalised summary of the relationships within orders of the families of seed

plants There are a few families that are not recognised even as options in APG II and

vice versa as well as a few extra orders The families to be recognised in the most

recent edition of Mabberleys The Plant Book (Mabberley 2008) are also largely

consistent with those below But all differences are trivial and will - I hope - eventually

disappear

Square brackets - [] - enclose clades the plus sign - + - designates sister taxa a

comma - - denotes part of a polytomy and quotation marks - - denotes a

paraphyletic group

SEED PLANTS

GYMNOSPERMS

Cycadales

Cycadaceae + Zamiaceae

Ginkgoales

Ginkgoaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

31

Pinales

Pinaceae [[Araucariaceae [Podocarpaceae + Phyllocladaceae]] [Sciadopityaceae

[Taxaceae + Cupressaceae]]]

Gnetales

Ephedraceae [Gnetaceae + Welwitschiaceae]

ANGIOSPERMSFLOWERING PLANTS

Amborellales

Amborellaceae

Nymphaeales

Hydatellaceae [Cabombaceae + Nymphaeaceae]

Austrobaileyales

Austrobaileyaceae [Schisandraceae + Trimeniaceae]

Chloranthales

Chloranthaceae

MAGNOLIIDS

Magnoliales

Myristicaceae [Magnoliaceae [[Himantandraceae + Degeneriaceae] [Eupomatiaceae +

Annonaceae]]]

Laurales

Calycanthaceae [[Siparunaceae [Gomortegaceae + Atherospermataceae]] [Monimiaceae

[Hernandiaceae + Lauraceae]]]

Canellales

Canellaceae + Winteraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

32

Piperales

[Hydnoraceae + Aristolochiaceae] [Piperaceae + Saururaceae]

MONOCOTS

Acorales

Acoraceae

Alismatales

Araceae [Tofieldiaceae [[Alismataceae [Hydrocharitaceae + Butomaceae]]

[Scheuchzeriaceae [Aponogetonaceae [Juncaginaceae [Maundiaceae [[Posidoniaceae

[Ruppiaceae + Cymodoceaceae]] [Zosteraceae + Potamogetonaceae]]]]]]]

Petrosaviales

Petrosaviaceae

Dioscoreales

Nartheciaceae [[Taccaceae + Thismiaceae] [Burmanniaceae + Dioscoreaceae]]

Pandanales

Velloziaceae Triuridaceae Stemonaceae [Pandanaceae + Cyclanthaceae]

Liliales

Corsiaceae [Campynemataceae [Petermanniaceae [Colchicaceae + Alstroemeriaceae]]

Melanthiaceae [[Philesiaceae + Rhipogonaceae] [Smilacaceae + Liliaceae]]]

Asparagales

Orchidaceae [[Boryaceae [Blandfordiaceae [Lanariaceae [Asteliaceae + Hypoxidaceae]]]]

[[Ixioliriaceae + Tecophilaeaceae] [Doryanthaceae [Iridaceae [Xeronemataceae

[Xanthorrhoeaceae [Alliaceae + Asparagaceae]]]]]]]

COMMELINIDS

Unplaced

Dasypogonaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

33

Arecales

Arecaceae

Poales

[Typhaceae + Bromeliaceae] [Rapateaceae [[Thurniaceae [Juncaceae + Cyperaceae]]

[[Mayacaceae [Eriocaulaceae + Xyridaceae]] [Flagellariaceae [Anarthriaceae

[Centrolepidaceae + Restionaceae]]] [Joinvilleaceae [Ecdeiocoleaceae + Poaceae]]]]]]]

Commelinales

[Commelinaceae + Hanguanaceae] [Philydraceae [Haemodoraceae + Pontederiaceae]]

Zingiberales

Musaceae [Strelitziaceae + Lowiaceae] Heliconiaceae [[Cannaceae + Marantaceae]

[Costaceae + Zingiberaceae]]

Ceratophyllales

Ceratophyllaceae

EUDICOTS

Ranunculales

Eupteleaceae [Papaveraceae [[[Lardizabalaceae + Circaeasteraceae] [Menispermaceae

[Berberidaceae + Ranunculaceae]]]]

Sabiales

Sabiaceae

Proteales

Nelumbonaceae [Platanaceae + Proteaceae]

Trochodendrales

Trochodendraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

34

Buxales

Haptanthaceae [Buxaceae + Didymelaceae]

CORE EUDICOTS

Gunnerales

Gunneraceae + Myrothamnaceae

Dilleniales

Dilleniaceae

Saxifragales

Peridiscaceae [[Paeoniaceae [Altingiaceae [Hamamelidaceae [Cercidiphyllaceae +

Daphniphyllaceae]]]] [[Crassulaceae [Aphanopetalaceae [Tetracarpaeaceae

[Penthoraceae + Haloragaceae]]]] [Iteaceae [Grossulariaceae + Saxifragaceae]]]]

Cynomoriaceae unplaced

Vitales

Vitaceae

ROSIDS

FABIDROSID I

Zygophyllales

Krameriaceae + Zygophyllaceae

Celastrales

Lepidobotryaceae + Celastraceae

Oxalidales

Huaceae [[Connaraceae + Oxalidaceae] [Cunoniaceae [Elaeocarpaceae [Brunelliaceae +

Cephalotaceae]]]]

Malpighiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

35

[Achariaceae [Goupiaceae [Violaceae + Passifloraceae] [Lacistemataceae Salicaceae]]

[Lophopyxidaceae Putranjivaceae] [Ctenolophonaceae [Erythroxylaceae +

Rhizophoraceae]] Linaceae Ixonanthaceae Humiriaceae Irvingiaceae

Centroplacaceae Pandaceae [[Bonnetiaceae + Clusiaceae] [Calophyllaceae

[Hypericaceae + Podostemaceae]]] [Malpighiaceae + Elatinaceae] Ochnaceae

[Peraceae [Rafflesiaceae + Euphorbiaceae]] [Phyllanthaceae + Picrodendraceae]

[Balanopaceae [[Trigoniaceae + Dichapetalaceae] [Chrysobalanaceae +

Euphroniaceae]]] Caryocaraceae

N-FIXING CLADE

Fabales

Quillajaceae [Fabaceae [Polygalaceae + Surianaceae]]

Rosales

Rosaceae [[Barbeyaceae [Elaeagnaceae Dirachmaceae Rhamnaceae]] [Ulmaceae

[Cannabaceae [Moraceae + Urticaceae]]]]

Cucurbitales

Anisophylleaceae [[Corynocarpaceae + Coriariaceae] [Cucurbitaceae [Tetramelaceae

[Datiscaceae + Begoniaceae]]]] Apodanthaceae

Fagales

Nothofagaceae [Fagaceae [[Myricaceae + Juglandaceae] [Casuarinaceae

[Ticodendraceae + Betulaceae]]]]

MALVIDROSID II

pgtGeraniales

Geraniaceae [[Melianthaceae + Francoaceae] [Vivianaceae + Ledocarpaceae]]

Myrtales

Combretaceae [[Onagraceae + Lythraceae] [[Vochysiaceae + Myrtaceae]

[Melastomataceae [Crypteroniaceae [Alzateaceae + Penaeaceae]]]]]

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

36

Crossosomatales

[Staphyleaceae [Guamatelaceae [Crossosomataceae + Stachyuraceae]]] [Aphloiaceae

[Geissolomataceae + Strasburgeriaceae]]

Picramniales

Picramniaceae

Sapindales

Biebersteiniaceae Nitrariaceae [Kirkiaceae [Anacardiaceae + Burseraceae]]

[Sapindaceae [Simaroubaceae Rutaceae Meliaceae]]

Huerteales

Gerradinaceae [Dipentodontaceae + Tapisciaceae]

Malvales

Neuradaceae [Thymelaeaceae [Sphaerosepalaceae Bixaceae [Cistaceae

[Sarcolaenaceae + Dipterocarpaceae]] [Cytinaceae + Muntingiaceae] Malvaceae]]

Brassicales

[Akaniaceae + Tropaeolaceae] [[Moringaceae + Caricaceae] [Setchellanthaceae

[Limnanthaceae [[Koeberliniaceae [Bataceae + Salvadoraceae]] [Emblingiaceae

[Pentadiplandraceae [Gyrostemonaceae Resedaceae Stixis and relatives] Tovariaceae

[Capparaceae [Cleomaceae + Brassicaceae]]]]]]]]

Berberidopsidales

Aextoxicaceae + Berberidopsidaceae

Santalales

Erythropalaceae Olacaceae [[Loranthaceae [Misodendraceae + Schoepfiaceae]]

[Opiliaceae + Santalaceae]] Balanophoraceae unplaced

Caryophyllales

[[Droseraceae [Nepenthaceae [Drosophyllaceae [Ancistrocladaceae +

Dioncophylleaceae]]]]] [[Frankeniaceae + Tamaricaceae] [Polygonaceae +

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

37

Plumbaginaceae]]] [Rhabdodendraceae [Simmondsiaceae [[Asteropeiaceae +

Physenaceae] [[Caryophyllaceae [Achatocarpaceae + Amaranthaceae]]

[Stegnospermataceae [Limeaceae [[Lophocarpaceae [Barbeuiaceae [Aizoaceae

[Sarcobataceae Phytolaccaceae Nyctaginaceae]]]] [Molluginaceae [Halophytaceae

Montiaceae Basellaceae Didiereaceae [Talinaceae [Portulacaceae [Anacampseros etc +

Cactaceae]]]]]]]]]]]]]

ASTERIDS

Cornales

[Cornaceae + Nyssaceae] [Hydrangeaceae + Loasaceae] Hydrostachyaceae [Curtisiaceae

+ Grubbiaceae]

Ericales

[Balsaminaceae [Marcgraviaceae + Tetrameristaceae]] [[Polemoniaceae +

Fouquieraceae] Lecythidaceae [[Sladeniaceae + Pentaphylacaceae] [Sapotaceae

[Ebenaceae [Maesaceae [Theophrastaceae [Myrsinaceae + Primulaceae]]]]]

[Mitrastemonaceae Theaceae [Symplocaceae [Styracaceae + Diapensiaceae]]

[[Sarraceniaceae [Roridulaceae + Actinidiaceae] [Clethraceae [Cyrillaceae +

Ericaceae]]]]]]

LAMIIDASTERID I

Unplaced

Metteniusaceae Oncothecaceae Icacinaceae and some unplaced ex-Icacinaceae

Garryales

Garryaceae + Eucommiaceae

Unplaced

Boraginaceae Vahliaceae

Gentianales

Rubiaceae [Gentianaceae [Loganiaceae [Gelsemiaceae + Apocynaceae]]]

Lamiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

38

Plocospermataceae [Oleaceae [Tetrachondraceae [Calceolariaceae

[Gesneriaceae Peltanthera [Pedaliaceae Carlemanniaceae Martyniaceae Stilbaceae

Plantaginaceae Linderniaceae Bignoniaceae Verbenaceae Lamiaceae Paulowniaceae

Schlegeliaceae Thomandersiaceae Phrymaceae Rehmannia Acanthaceae

Scrophulariaceae Orobanchaceae Byblidaceae Lentibulariaceae]]]]]

Solanales

[Montiniaceae [Sphenocleaceae + Hydroleaceae]] [Convolvulaceae + Solanaceae]

CAMPANULIDASTERID II

Aquifoliales

[[Cardiopteridaceae + Stemonuraceae] [Aquifoliaceae [Helwingiaceae +

Phyllonomaceae]]

Asterales

[Rousseaceae + Campanulaceae] Pentaphragmataceae [[Alseuosmiaceae [Phellinaceae

+ Argophyllaceae]] Stylidiaceae [Menyanthaceae [Goodeniaceae [Calyceraceae +

Asteraceae]]]]

Escalloniales

Escalloniaceae

Bruniales

[Bruniaceae + Columelliaceae]

Apiales

Pennantiaceae [Torricelliaceae [Griseliniaceae [Pittosporaceae [Araliaceae

[Myodocarpaceae + Apiaceae]]]]]

Paracryphiales

Paracryphiaceae

Dipsacales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

39

Adoxaceae [Diervillaceae [Caprifoliaceae [Linnaeaceae [Morinaceae [Dipsacaceae +

Valerianaceae]]]]]

LINKS TO ORDERS AND FAMILIES

[Back to Top]

MAIN GROUPINGS

Asterids commelinids core eudicots asterid 1 asterid

2 eudicots gymnosperms Magnoliophyta monocots N-fixing claderosids seed plants

ALPHABETICAL LISTING OF ALL ORDINAL NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | L | M | N | O | P | Q | R | S | T | U | V | W | X | Z

Abietales Acanthales Acerales Acorales Actinidiales Actinostrobales Adoxales Aescul

ales Agavales Aizoales AkanialesAlismatales Alliales Alseuosmiales Alstroemeriales

Altingiales Amaranthales Amaryllidales Amborellales AmbrosialesAmmiales Amomal

es Ancistrocladales Anisophylleales Annonales Anthobolales Apiales Apocynales Apo

nogetonalesAquifoliales Arales Araliales Aralidiales Araucariales Arecales Aristolochi

ales Athrotaxidales Asarales AsclepiadalesAsparagales Asphodelales Asparagales Ast

eliales Atriplicales Aucubales Austrobaileyales Avenales

Balanitales Balanopales Balanophorales Balsaminales Barbeyales Barclayales Batales

Begoniales BerberidalesBerberidopsidales Betulales Biebersteiniales Bignoniales Bix

ales Boraginales Brassicales Brexiales Bromeliales BrunialesBrunoniales Burmannial

es Burserales Butomales Buxales Byblidales

Cactales Callitrichales Calycanthales Calycerales Campanulales Campynematales Can

ellales Cannales CapparalesCaprifoliales Cardiopteridales Carduales Caricales Carle

manniales Caryophyllales Cassiales Casuarinales CelastralesCentrolepidales Cephalot

ales Cephalotaxales Ceratophyllales Cercidiphyllales Chenopodiales Chironiales Chlor

anthalesChrysobalanales Cinchonales Circaeasterales Cistales Citrales Cocosales Col

chicales Columelliales CombretalesCommelinales Connarales Convolvulales Coriarial

es Cornales Corylales Corynocarpales Crassulales CrossosomatalesCucurbitales Cunn

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

40

inghamiales Cunoniales Cupressales Cycadales Cyclanthales Cymodoceales Cynarales

CynomorialesCyperales Cyrillales Cytinales

Daphnales Daphniphyllales Dasypogonales Datiscales Degeneriales Desfontainiales D

iapensiales Didymelales DillenialesDioales Dioncophyllales Dioscoreales Diospyrales

Dipentodontales Dipsacales Droserales

Ebenales Echiales Elaeagnales Elaeocarpales Elatinales Elodeales Emmotales Empetr

ales Ephedrales Ericales EriocaulalesErythropalaless Escalloniales Eucommiales Euph

orbiales Eupomatiales Eupteleales Euryalaless

Fabales Fagales Falcatifoliales Ficales Flacourtiales Flagellariales Fouquieriales Franc

oales Frangulales

Galiales Garryales Geissolomatales Gentianales Geraniales Gesneriales Ginkgoales G

laucidiales Globulariales GnetalesGoodeniales Greyiales Griseliniales Grossulariales

Grubbiales Gunnerales Gyrocarpales Gyrostemonales

Haemodorales Haloragales Hamamelidales Hanguanales Heisteriales Helleborales He

lwingiales Himantandrales HippuridalesHomaliales Hortensiales Huales Huerteales

Hydatellales Hydnorales Hydrangeales Hydrastidales HydrocharitalesHydropeltidales

Hydrostachyales Hypericales Hypoxidales

Icacinales Illiciales Iridales Irvingales Iteales Ixiales

Jasminales Juglandales Julianiales Juncaginales Juncales

Lacistematales Lactoridales Lamiales Lardizabalales Laurales Lecythidales Ledocarpal

es Leitneriales LentibularialesLigustrales Liliales Limnanthales Linales Liriales Loasal

es Lobeliales Loganiales Lonicerales Loranthales LowialesLythrales

Magnoliales Malpighiales Malvales Marathrales Marcgraviales Mayacales Medusagy

nales Medusandrales MelanthialesMelastomatales Meliales Melianthales Meliosmal

es Menispermales Menyanthales Metteniusales MitrastemonalesMiyoshiales Moni

miales Moringales Musales Myricales Myristicales Myrothamnales Myrsinales Myrta

les

Najadales Nandinales Narcissales Nartheciales Nelumbonales Nepenthales Neuradal

es Nitrariales Nolanales NothofagalesNyctaginales Nymphaeales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

41

Ochnales Oenotherales Olacales Oleales Onagrales Oncothecales Opuntiales Orchid

ales Oxalidales

Saxifragales Pandales Pandanales Papaverales Paracryphiales Parasitaxales Paridales

Parnassiales PassifloralesPenaeales Pennantiales Pentaphragmatales Peridiscales P

etiveriales Petrosaviales Phellinales Philydrales PhyllanthalesPhysenales Phytolaccal

es Picramniales Pinales Pinguiculales Piperales Pittosporales Plantaginales Platanale

sPlumbaginales Poales Podocarpales Podophyllales Podostemales Polemoniales Pol

ygalales Polygonales PontederialesPortulacales Posidoniales Potamogetonales Primu

lales Proteales

Quercales Quillajales Quintiniales

Rafflesiales Ranunculales Rapateales Resedales Restionales Rhabdodendrales Rhamn

ales Rhinanthales RhizophoralesRhodorales Rhoipteleales Roridulales Rosales Rouss

eales Rubiales Ruppiales Rutales

Sabiales Salicales Salvadorales Samolales Samydales Sanguisorbales Santalales Sapin

dales Sapotales SarracenialesSaxegotheales Saxifragales Scheuchzeriales Sciadopitya

les Scleranthales Scrophulariales Scyphostegiales SedalesSilenales Simmondsiales S

milacales Solanales Sphenocleales Sphenostemonales Stangeriales Stellariales Stemo

nalesStilbales Stylidiales Styracales Surianales

Taccales Tamales Tamaricales Taxales Taxodiales Tecophilaeales Terebinthales Tern

stroemiales Theales TheligonalesThymelaeales Tiliales Tofieldiales Torricelliales Tov

ariales Tribelales Trilliales Trimeniales Triuridales TrochodendralesTropaeolales Turn

erales Typhales

Ulmales Urticales

Vacciniales Vahliales Vallisneriales Velloziales Veratrales Verbenales Viburnales Vinc

ales Violales Viscales VitalesVochysiales

Welwitschiales Winterales

Xanthorrhoeales Ximeniales Xyridales

Zamiales Zingiberales Zosterales Zygophyllales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

42

ALPHABETICAL LISTING OF ALL FAMILY NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | K | L | M | N | O | P | Q | R | S | T | U | V | W | X | Y

|Z

Back to Home

In some cases clicking on a family synonym will send you directly to the subfamily in

which the genus of the basionym is to be found

Abietaceae Abolbodaceae Abrophyllaceae Acalyphaceae Acanthaceae [Acanthochla

mydaceae] Acarnaceaee AceraceaeAchariaceae Achatocarpaceae Achradaceae Achy

ranthaceae Acmopylaceae Acoraceae Acristaceae ActaeaceaeActinidiaceae Actinota

ceae Amentotaxaceae Adenogrammaceae Adoxaceae Aegialitidaceae Aegicerataceae

Aegilopaceae Aeginetiaceaee Aegiphilaceae Aesculaceae Aextoxicaceae Agapanthac

eae Agavaceae AgdestidaceaeAgialidaceae Agrimoniaceae Agrostidaceae Ailanthace

ae Aitoniaceae Aizoaceae Akaniaceae Alangiaceae AlchemillaceaeAldrovandaceae A

lismataceae Alliaceae Allioniaceae Allophylaceae Aloaceae Alopecuraceae Alpiniacea

e AlseuosmiaceaeAlsinaceae Alsodeiaceae Alstroemeriaceae Altingiaceae Alzateacea

e Amaranthaceae Amaryllidaceae Amborellaceae Ambrosiaceae Amentotaxaceae A

mmanniaceae Ammiaceae Amomaceae Ampelidopsaceae Amygdalaceae Amyridacea

eAnacardiaceae Anagallidaceae Anarthriaceae Anchusaceae Ancistrocladaceae Andr

omedaceae AndropogonaceaeAndrostachyaceae Androsynaceae Anemarrhenaceae

Anemonaceae Angelicaceae Anisophylleaceae AnnonaceaeAnomochloaceae Anopter

aceae Anrederaceae Anthemidaceae Anthericaceae Anthobolaceae Antidesmataceae

Antirrhinaceae Antoniaceae Aparinaceae Aphanopetalaceae Aphloiaceae Aphyllanth

aceae Apiaceae ApocynaceaeApodanthaceae Aponogetonaceae Aporusaceae Apose

ridaceae Apostasiaceae Aptandraceae AquifoliaceaeAquilariaceae Aquilegiaceae A

raceae Aragoaceae Araliaceae Aralidiaceaee Araucariaceae Arbutaceae Arceuthidace

aeArceuthobiaceae Arctostaphylaceae Arctotidaceae Ardisiaceae Arecaceae Argoph

yllaceae Arisaraceae AristolochiaceaeAristoteliaceae Arjonaceae Armeriaceae Artem

isiaceae Arthrotaxidaceae Artocarpaceae Arundinaceae ArundinellaceaeAsaraceae A

sclepiadaceae Ascyraceaee Asiraceae Asparagaceae Asperulaceae Asphodelaceae As

pidistraceaeeAsteliaceae Asteraceae Asteranthaceae Asteropeiaceae Astragalaceae

Astrocarpaceae AthanasiaceaeAtherospermataceae Atriplicaceae Atropaceae Aucuba

ceae Aurantiaceae Austrobaileyaceae Austrotaxaceae AvenaceaeAverrhoaceae Avet

raceae Avicenniaceae Azaleaceae Azimaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

43

Balanitaceae Balanopaceae Balanophoraceae Balsameaceae Balsaminaceae Bambusa

ceae Barbaceniaceae BarbeuiaceaeBarbeyaceae Barclayaceae Barringtoniaceae Base

llaceae Bataceae Baueraceae Bauhiniaceae Baxteriaceae BegoniaceaeBehniaceae B

elangeraceae Belloniaceae Bembiciaceae Berberidaceae Berberidopsidaceae Berryac

eaee BersamaceaeBertyaceae Berzeliaceae Besleriaceae Betaceae Betulaceae Biebe

rsteiniaceae Bifariaceae Bignoniaceae BischofiaceaeBixaceae Blakeaceae Blandfordia

ceae Blattiaceae Blepharocaryaceae Blitaceae Blyxaceae Boerlagellaceae Bolivaracea

eBombacaceae Bonnetiaceae Bontiaceae Boopidaceae Boraginaceae Borassaceae B

oroniaceae BoryaceaeBotryodendraceae Bougainvilleaceae Boweniaceae Brassicacea

e Brachycaulaceae Bretschneideraceae BrexiaceaeBromeliaceae Brownlowiaceae Br

unelliaceae Bruniaceae Brunoniaceae Brunsvigiaceae Bryoniaceae BuchneraceaeBuci

daceae Buddlejaceae Buglossaceae Bulbocodiaceae Bumeliaceae Bupleuraceae Burc

hardiaceae BurmanniaceaeBurseraceae Butneriaceae Butomaceae Buxaceae Byblida

ceae Byttneriaceae

Cabombaceae Cacaoaceae Cactaceae Caesalpiniaceae Caladiaceae Calamaceae Calce

olariaceae CalectasiaceaeCalendulaceae Callaceae Callicomaceae Calligonaceae Calli

traceae Callitrichaceae Calochortaceae CalophyllaceaeCalthaceae Calycanthaceae Ca

lyceraceae Cambogiaceae Camelliaceae Campanulaceae CampynemataceaeCanacom

yricaceae Canellaceae Cannabaceae Cannaceae Canopodaceae Canotiaceae Cansjera

ceae CapparaceaeCaprariaceae Caprifoliaceae Cardiopteridaceae Carduaceae Carica

ceae Carissaceae Carlemanniaceae CarpinaceaeCarpodetaceae Cartonemataceae Ca

ryocaraceae Caryophyllaceae Casearieaceae Cassiaceae CassipoureaceaeCassythacea

e Castaneaceae Castelaceae Casuarinaceae Catesbaeaceae Cathedraceae Cecropiace

ae CedraceaeCedrelaceae Celastraceae Celosiaceae Celtidaceae Centaureaceae Cen

trolepidaceae Centroplacaceae CepaceaeCephalanthaceae Cephalotaceae Cephalota

xaceae Cerastiaceae Ceratoniaceae Ceratophyllaceae CerberaceaeCercidiphyllaceae

Cercocarpaceae Cercodiaceae Cereaceae Cerinthaceae Ceroxylaceae Cestraceae Cev

alliaceaeChailletiaceae Chamaedoraceae Chamaemoraceae Chamelauciaceae Chaun

ochitonaceae Chelidoniaceae ChelonaceaeChenopodiaceae Chimonanthaceae Chingi

thamnaceae Chionographidaceae Chiranthodendraceae ChironiaceaeChlaenaceae Ch

loanthaceae Chloranthaceae Chloridaceae Chlorogalaceae Chrysobalanaceae Cicerac

eae CichoriaceaeCimicifugaceae Cinchoniaceae Circaeaceae Circaeasteraceae Cissac

eae Cistaceae Citraceae Clematidaceae CleomaceaeClethraceae Clusiaceae Cneorac

eae Cnestidaceae Cnicaceae Cobaeaceae Cochlospermaceae Cocosaceae Coffeaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

44

Colchicaceae Coleogynaceae Columelliaceae Combretaceae Commelinaceae Comocl

adiaceae Compositae CompsoaceaeConantheraceae Connaraceae Conostylidaceae C

onvallariaceae Convolvulaceae Cordiaceae CoreopsidaceaeCoriandraceae Coriariacea

e Coridaceae Corispermaceae Cornaceae Corokiaceae Coronillaceae Correaceae Cor

rigiolaceaeCorsiaceae Corydalaceae Corylaceae Corynocarpaceae Coryphaceae Cost

aceae Cotyledonaceae CoulaceaeCoutariaceae Coutoubiaceae Crassulaceae Crescen

tiaceae Cressaceae Crinaceae Crocaceae CroomiaceaeCrossosomataceae Crotonacea

e Cruciferae Cryptaceae Crypteroniaceae Cryptocorynaceae CryptomeriaceaeCtenol

ophonaceae Cucurbitaceae Cunninghamiaceae Cunoniaceae Cupressaceae Curcumac

eae Curtisiaceae CuscutaceaeCyananthaceae Cyanastraceae Cynanchaceae Cyanella

ceae Cycadaceae Cyclanthaceae CyclantheraceaeCyclocheilaceae Cydoniaceae Cymo

doceaceae Cynaraceae Cynocrambaceae Cynomoriaceae Cyperaceae CyphiaceaeCyp

hocarpaceae Cypripediaceae Cyrillaceae Cyrtandraceae Cyrtanthaceae Cytinaceae

Dacrycarpaceae Dactylanthaceae Damasoniaceae Daphnaceae Daphniphyllaceae Das

ypogonaceae DatiscaceaeDaturaceae Daucaceae Davidiaceae Davidsoniaceae Decais

neaceae Deeringiaceae Degeneriaceae DelphiniaceaeDendrophthoaceae Columelliac

eae Detariaceae Dialypetalanthaceae Dianellaceae Dianthaceae DiapensiaceaeDicha

petalaceae Dichondraceae Diclidantheraceae Dicrastylidiaceae Dictamnaceae Didiere

aceae DidymelaceaeDidymocarpaceae Diegodendraceae Diervillaceae Digitalidaceae

e Dilatridaceae Dilleniaceae Dioaceae DionaeaceaeDioncophyllaceae Dioscoreaceae

Diosmaceae Diospyraceae Dipentodontaceae Diphylleiaceae Diplolaenaceae Dipsaca

ceaeDipterocarpaceae Dirachmaceae Disanthaceae Diselmaceae Dodonaeaceae Do

mbeyaceae Donatiaceae DorsteniaceaeDortmannaceae Doryanthaceae Drabaceae D

racaenaceae Dracontiaceae Drimyidaceae Droseraceae DrosophyllaceaeDryadaceae

Duabangaceae Duckeodendraceae Dulongiaceae Durantaceae Durionaceae Dysphani

aceae

Ebenaceae Eccremidaceae Ecdeiocoleaceae Echiaceae Echinopaceae Ehretiaceae Ela

eagnaceae ElaeocarpaceaeElegiaceae Elatinaceae Ellisiaceae Ellisiophyllaceae Elodea

ceae Elytranthaceae Embeliaceae Emblingiaceae EmmotaceaeEmpetraceae Encepha

lartaceae Engelhardtiaceae Enhalaceae Epacridaceae Ephedraceae Ephemeranthacea

eEpilobiaceae Epimediaceae Eragrostidaceae Eremolepidaceae Eremosynaceae Erica

ceae Erinaceae EriocaulaceaeEriogonaceae Eriospermaceae Erodiaceae Erycibaceae

Eryngiaceae Erysimaceae Erythroniaceae ErythropalaceaeErythrospermaceae Erythro

xylaceae Escalloniaceae Eschscholziaceae Eucomidaceae Eucommiaceae Eucryphiace

aeEuonymaceae Eupatoriaceae Euphorbiaceae Euphrasiaceae Euphroniaceae Eupo

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

45

matiaceae Eupteleaceae EuryalaceaeEustrephaceae Euthemidaceae Eutocaceae Exb

ucklandiaceae Exocarpaceae

Fabaceae Fagaceae Falcatifoliaceae Ferulaceae Festucaceae Ficaceae Ficoidaceae Fit

zroyaceae FlacourtiaceaeFlagellariaceae Flindersiaceae Foetidiaceae Forestieraceae

Fothergillaceae Fouquieriaceae Fragariaceae FrancoaceaeFrangulaceae Frankeniacea

e Fraxinaceae Fraxinellaceae Freycinetiaceae Fritillariaceae Fuchsiaceae Fumariacea

eFunkiaceae

Gaiadendraceae Galacaceae Galanthaceae Galaxiaceae Galedupaceae Galeniaceae G

aliaceae Garciniaceae GardeniaceaeGardneriaceae Garryaceae Geissolomataceae Ge

itonoplesiaceae Gelsemiaceae Geniostomaceae GentianaceaeGeonomataceae Geosir

idaceae Geraniaceae Gerrardinaceae Gesneriaceae Gethyllidaceae Gilliesiaceae Gina

lloaceaeGinkgoaceae Gisekiaceae Gladiolaceae Glaucidiaceae Glechomaceae Glinace

ae Globulariaceae Gnaphaliaceae GnetaceaeGoetzeaceae Gomortegaceae Gomphiac

eae Gomphrenaceae Gonystylaceae Goodeniaceae Gordoniaceae GouaniaceaeGoup

iaceae Gramineae Gratiolaceae Grewiaceae Greyiaceae Grielaceae Griseliniaceae Gr

onoviaceae GrossulariaceaeGrubbiaceae Guaiacanaceae Guamatelaceae Guettardac

eae Gunneraceae Gustaviaceae Guttiferae GyrocarpaceaeGyrostemonaceae

Hachetteaceae Haemanthaceae Haemodoraceae Halesiaceae Hallieraceae Halocarpa

ceae Halophilaceae HalophytaceaeHaloragaceae Hamamelidaceae Hanguanaceae H

aptanthaceae Harmandaceae Hebenstretiaceae HectorellaceaeHederaceae Hedyosm

aceae Hedyotidaceae Hedysaraceae Heisteriaceae Heleniaceae Heliamphoraceae H

elianthaceaeHelianthemaceae Helichrysaceae Heliconiaceae Helicteraceae Heliotropi

aceae Helleboraceae Heloniadaceae HelosidaceaeHelwingiaceae Hemerocallidaceae

Hemimeridaceae Hemiphylacaceae Henriqueziaceae Henslowiaceae Hermanniaceae

Hernandiaceae Herniariaceae Herreriaceae Hesperocallidaceae Heterantheraceae He

teropyxidaceae HeterostylaceaeHibbertiaceae Hibiscaceae Hilleriaceae Himantandra

ceae Hippocastanaceae Hippocrateaceae HippomanaceaeHippophaeaceae Hippurida

ceae Hirtellaceae Holacanthaceae Homaliaceae Hopkinsiaceae Hoplestigmataceae H

ordeaceaeHornschuchiaceae Hortensiaceae Hortoniaceae Hostaceae Houstoniaceae

Huaceae Huerteaceae HugoniaceaeHumbertiaceae Humiriaceae Hyacinthaceae Hyd

atellaceae Hydnoraceae Hydrangeaceae Hydrastidaceae HydrillaceaeHydrocharitacea

e Hydrocotylaceae Hydrogetonaceae Hydroleaceae Hydropeltidaceae Hydrophylacac

eae HydrophyllaceaeHydrostachyaceae Hymenocardiaceae Hypecoaceae Hyperanth

eraceae Hypericaceae Hypopityaceae HypoxidaceaeHypseocharitaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

46

Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta

ceae ImperatoriaceaeInocarpaceae Inulaceae Ionidiaceae Iriarteaceae Iridaceae Irvi

ngiaceae Isnardiaceae Isophysidaceae IteaceaeStrasburgeriaceae Ixiaceae Ixioliriace

ae Ixonanthaceae

Jalapaceae Jamboliferaceae Japonoliriaceae Jasionaceae Jasminiaceae Johnsoniaceae

Joinvilleaceae JuglandaceaeJulianiaceae Juncaceae Juncaginaceae Juniperaceae Jus

siaeaceae Justiciaceae

Kadsuraceae Kaliphoraceae Kalmiaceae Kaniaceae Kiggelariaceae Kingdoniaceae Kin

giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae

Krameriaceae

Labiatae Lacandoniaceae Lachenaliaceae Lacistemataceae Lactoridaceae Lactucaceae

Lagerstroemiaceaee LamiaceaeLampsanaceae Lanariaceae Langsdorffiaceae Lantan

aceae Lapageriaceae Lardizabalaceae Lasiopetalaceae LauraceaeLawsoniaceae Laxm

anniaceae Lecythidaceae Ledaceae Ledocarpaceae Leeaceae Leguminosae Leitneriac

eae LemnaceaeLennoaceae Lentibulariaceae Lentiscaceae Leoniaceae Leonticaceae

Lepidobotryaceae LepidocarpaceaeLepidocaryaceae Lepidocerataceae Leptaulaceae

Leptospermaceae Lepturaceae Lepuropetalaceae LeuctenbergiaceaeLibocedraceae

Licaniaceae Ligustraceae Lilaeaceae Liliaceae Limeaceae Limnanthaceae Limnocharit

aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace

ae Linderniaceae Lindleyaceae Linnaeaceae LippayaceaeLiriaceae Liriodendraceae Li

ssocarpaceae Littorellaceae Loasaceae Lobeliaceaee Loganiaceae LomandraceaeLoni

ceraceae Lopeziaceae Lophiocarpaceae Lophiolaceae Lophiraceae Lophophytaceae L

ophopyxidaceae LoranthaceaeLotaceae Lowiaceae Lupulaceaee Luxemburgiaceae Lu

zuriagaceae Lychnidaceae Lyciaceae LyginiaceaeLygodisodeaceae Lysimachiaceae Ly

thraceae

Macarisiaceae Mackinlayaceae Madiaceae Maesaceae Magnoliaceae Malaceae Male

sherbiaceae MalortieaceaeMalpighiaceae Malvaceae Mangiaceae Manicariaceae Ma

paniaceae Marantaceae Marathraceae MarcgraviaceaeMartyniaceae Mastixiaceae

Matricariaceae Maundiaceae Mayacaceae Medeolaceae Medusagynaceae Peridiscac

eaeMelaleucaceae Melampyraceae Melanophyllaceae Melanthiaceae Melastomatac

eae Meliaceae Melianthaceae MelicaceaeMeliosmaceae Melittidaceae Melochiacea

e Memecylaceae Mendonciaceae Menispermaceae Menthaceae MenyanthaceaeMe

nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

47

Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

Nypaceae Nyssaceae

Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

ceae Papilionaceae Pappophoraceae PapyraceaeParacryphiaceae Parasitaxaceae Par

ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

48

hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

49

cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

50

ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 29: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

29

metazoans However Pax 6 seems to be a master control gene that is involved in eye

formation perhaps throughout all bilateralians (eg Gehring amp Ikeo 1999 Erwin amp

Davidson 2002) which changes the nature of the problem Note however that in

general our invocation of evolutionary tendencies to explain the patterns of

distributions of characters is really an admission that we do not understand those

patterns

As we find out more about variation we will probably find fewer and fewer features

constant throughout a group Most unqualified statements of presence and absence

should properly be qualified as usually present or usually absent if one is thinking of

the characterisations on the order pages as encompassing the total variation within a

clade Thus Pistia alone among monocots as so far known has sieve tube plastids with

starch grains not protein crystals However this does not affect the fact that sieve tube

plastids with cuneate protein crystals and lacking starch grains are an apomorphy for

monocots Furthermore fossils in a number of cases suggest character combinations

unknown in extant taxa as may be seen in the discussions of Fagaceae Platanaceae

Iteaceae Calycanthaceae etc Confusing the issue there may be questions as to where

exactly on the tree a particular fossil is to be placed (eg see Nymphaeaceae

Calycanthaceae Archaefructus etc)

To summarize given our current understandings of both phylogenies and

characters evolution of some characters in which we are interested seems very labile

(see eg D Soltis et al 2005b Endress and Matthews 2006a Stevens 2006b Ekman et

al 2008 for a good recent study) and I have been cautious when talking about character

evolution Much effort must continue to be spent in summarizing characters of well-

established clades at all levels providing features by which they may be recognized and

signaling synapomorphies Remember that (1) the basic morphological anatomical and

chemical knowledge of many critical taxa is woefully incomplete (2) different

assumptions about character evolution may greatly affect the position of

synapomorphies on trees (3) in many cases relationships within and between many

groups are too uncertain at present to worry very much about synapomorphies and

(4) we must be clear about what we do and do not not know As mentioned above I

have indicated in a number of places where there are particularly important

uncertainties as to where characters should be placed on the tree Nevertheless it is a

relatively easy matter to update notes such as these and it can be a simple matter to

incorporate new data on characters that have never before been considered in the

context of a tree Much basic - and unfortunately perhaps unfashionable - work must be

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

30

carried to clarify the distribution of morphological anatomical and chemical characters

for examples of what can to be done see the work of P K Endress and collaborators

(floral morphology and development) and S R Jensen and collaborators (iridoids) But

acquiring information about nodal anatomy cell and tissue distribution nodal anatomy

and the like is not difficult at all and should be generally encouraged

But all this is merely a necesary prelude to the understanding of evolution For this

one needs to know a lot more including dating the phylogeny understanding the

function(s) of characters factoring in relevant aspects of the palaeoenvironment in

which an apomorphy first appeared etc

SUMMARY OF THE SYSTEM

Below is a formalised summary of the relationships within orders of the families of seed

plants There are a few families that are not recognised even as options in APG II and

vice versa as well as a few extra orders The families to be recognised in the most

recent edition of Mabberleys The Plant Book (Mabberley 2008) are also largely

consistent with those below But all differences are trivial and will - I hope - eventually

disappear

Square brackets - [] - enclose clades the plus sign - + - designates sister taxa a

comma - - denotes part of a polytomy and quotation marks - - denotes a

paraphyletic group

SEED PLANTS

GYMNOSPERMS

Cycadales

Cycadaceae + Zamiaceae

Ginkgoales

Ginkgoaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

31

Pinales

Pinaceae [[Araucariaceae [Podocarpaceae + Phyllocladaceae]] [Sciadopityaceae

[Taxaceae + Cupressaceae]]]

Gnetales

Ephedraceae [Gnetaceae + Welwitschiaceae]

ANGIOSPERMSFLOWERING PLANTS

Amborellales

Amborellaceae

Nymphaeales

Hydatellaceae [Cabombaceae + Nymphaeaceae]

Austrobaileyales

Austrobaileyaceae [Schisandraceae + Trimeniaceae]

Chloranthales

Chloranthaceae

MAGNOLIIDS

Magnoliales

Myristicaceae [Magnoliaceae [[Himantandraceae + Degeneriaceae] [Eupomatiaceae +

Annonaceae]]]

Laurales

Calycanthaceae [[Siparunaceae [Gomortegaceae + Atherospermataceae]] [Monimiaceae

[Hernandiaceae + Lauraceae]]]

Canellales

Canellaceae + Winteraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

32

Piperales

[Hydnoraceae + Aristolochiaceae] [Piperaceae + Saururaceae]

MONOCOTS

Acorales

Acoraceae

Alismatales

Araceae [Tofieldiaceae [[Alismataceae [Hydrocharitaceae + Butomaceae]]

[Scheuchzeriaceae [Aponogetonaceae [Juncaginaceae [Maundiaceae [[Posidoniaceae

[Ruppiaceae + Cymodoceaceae]] [Zosteraceae + Potamogetonaceae]]]]]]]

Petrosaviales

Petrosaviaceae

Dioscoreales

Nartheciaceae [[Taccaceae + Thismiaceae] [Burmanniaceae + Dioscoreaceae]]

Pandanales

Velloziaceae Triuridaceae Stemonaceae [Pandanaceae + Cyclanthaceae]

Liliales

Corsiaceae [Campynemataceae [Petermanniaceae [Colchicaceae + Alstroemeriaceae]]

Melanthiaceae [[Philesiaceae + Rhipogonaceae] [Smilacaceae + Liliaceae]]]

Asparagales

Orchidaceae [[Boryaceae [Blandfordiaceae [Lanariaceae [Asteliaceae + Hypoxidaceae]]]]

[[Ixioliriaceae + Tecophilaeaceae] [Doryanthaceae [Iridaceae [Xeronemataceae

[Xanthorrhoeaceae [Alliaceae + Asparagaceae]]]]]]]

COMMELINIDS

Unplaced

Dasypogonaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

33

Arecales

Arecaceae

Poales

[Typhaceae + Bromeliaceae] [Rapateaceae [[Thurniaceae [Juncaceae + Cyperaceae]]

[[Mayacaceae [Eriocaulaceae + Xyridaceae]] [Flagellariaceae [Anarthriaceae

[Centrolepidaceae + Restionaceae]]] [Joinvilleaceae [Ecdeiocoleaceae + Poaceae]]]]]]]

Commelinales

[Commelinaceae + Hanguanaceae] [Philydraceae [Haemodoraceae + Pontederiaceae]]

Zingiberales

Musaceae [Strelitziaceae + Lowiaceae] Heliconiaceae [[Cannaceae + Marantaceae]

[Costaceae + Zingiberaceae]]

Ceratophyllales

Ceratophyllaceae

EUDICOTS

Ranunculales

Eupteleaceae [Papaveraceae [[[Lardizabalaceae + Circaeasteraceae] [Menispermaceae

[Berberidaceae + Ranunculaceae]]]]

Sabiales

Sabiaceae

Proteales

Nelumbonaceae [Platanaceae + Proteaceae]

Trochodendrales

Trochodendraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

34

Buxales

Haptanthaceae [Buxaceae + Didymelaceae]

CORE EUDICOTS

Gunnerales

Gunneraceae + Myrothamnaceae

Dilleniales

Dilleniaceae

Saxifragales

Peridiscaceae [[Paeoniaceae [Altingiaceae [Hamamelidaceae [Cercidiphyllaceae +

Daphniphyllaceae]]]] [[Crassulaceae [Aphanopetalaceae [Tetracarpaeaceae

[Penthoraceae + Haloragaceae]]]] [Iteaceae [Grossulariaceae + Saxifragaceae]]]]

Cynomoriaceae unplaced

Vitales

Vitaceae

ROSIDS

FABIDROSID I

Zygophyllales

Krameriaceae + Zygophyllaceae

Celastrales

Lepidobotryaceae + Celastraceae

Oxalidales

Huaceae [[Connaraceae + Oxalidaceae] [Cunoniaceae [Elaeocarpaceae [Brunelliaceae +

Cephalotaceae]]]]

Malpighiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

35

[Achariaceae [Goupiaceae [Violaceae + Passifloraceae] [Lacistemataceae Salicaceae]]

[Lophopyxidaceae Putranjivaceae] [Ctenolophonaceae [Erythroxylaceae +

Rhizophoraceae]] Linaceae Ixonanthaceae Humiriaceae Irvingiaceae

Centroplacaceae Pandaceae [[Bonnetiaceae + Clusiaceae] [Calophyllaceae

[Hypericaceae + Podostemaceae]]] [Malpighiaceae + Elatinaceae] Ochnaceae

[Peraceae [Rafflesiaceae + Euphorbiaceae]] [Phyllanthaceae + Picrodendraceae]

[Balanopaceae [[Trigoniaceae + Dichapetalaceae] [Chrysobalanaceae +

Euphroniaceae]]] Caryocaraceae

N-FIXING CLADE

Fabales

Quillajaceae [Fabaceae [Polygalaceae + Surianaceae]]

Rosales

Rosaceae [[Barbeyaceae [Elaeagnaceae Dirachmaceae Rhamnaceae]] [Ulmaceae

[Cannabaceae [Moraceae + Urticaceae]]]]

Cucurbitales

Anisophylleaceae [[Corynocarpaceae + Coriariaceae] [Cucurbitaceae [Tetramelaceae

[Datiscaceae + Begoniaceae]]]] Apodanthaceae

Fagales

Nothofagaceae [Fagaceae [[Myricaceae + Juglandaceae] [Casuarinaceae

[Ticodendraceae + Betulaceae]]]]

MALVIDROSID II

pgtGeraniales

Geraniaceae [[Melianthaceae + Francoaceae] [Vivianaceae + Ledocarpaceae]]

Myrtales

Combretaceae [[Onagraceae + Lythraceae] [[Vochysiaceae + Myrtaceae]

[Melastomataceae [Crypteroniaceae [Alzateaceae + Penaeaceae]]]]]

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

36

Crossosomatales

[Staphyleaceae [Guamatelaceae [Crossosomataceae + Stachyuraceae]]] [Aphloiaceae

[Geissolomataceae + Strasburgeriaceae]]

Picramniales

Picramniaceae

Sapindales

Biebersteiniaceae Nitrariaceae [Kirkiaceae [Anacardiaceae + Burseraceae]]

[Sapindaceae [Simaroubaceae Rutaceae Meliaceae]]

Huerteales

Gerradinaceae [Dipentodontaceae + Tapisciaceae]

Malvales

Neuradaceae [Thymelaeaceae [Sphaerosepalaceae Bixaceae [Cistaceae

[Sarcolaenaceae + Dipterocarpaceae]] [Cytinaceae + Muntingiaceae] Malvaceae]]

Brassicales

[Akaniaceae + Tropaeolaceae] [[Moringaceae + Caricaceae] [Setchellanthaceae

[Limnanthaceae [[Koeberliniaceae [Bataceae + Salvadoraceae]] [Emblingiaceae

[Pentadiplandraceae [Gyrostemonaceae Resedaceae Stixis and relatives] Tovariaceae

[Capparaceae [Cleomaceae + Brassicaceae]]]]]]]]

Berberidopsidales

Aextoxicaceae + Berberidopsidaceae

Santalales

Erythropalaceae Olacaceae [[Loranthaceae [Misodendraceae + Schoepfiaceae]]

[Opiliaceae + Santalaceae]] Balanophoraceae unplaced

Caryophyllales

[[Droseraceae [Nepenthaceae [Drosophyllaceae [Ancistrocladaceae +

Dioncophylleaceae]]]]] [[Frankeniaceae + Tamaricaceae] [Polygonaceae +

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

37

Plumbaginaceae]]] [Rhabdodendraceae [Simmondsiaceae [[Asteropeiaceae +

Physenaceae] [[Caryophyllaceae [Achatocarpaceae + Amaranthaceae]]

[Stegnospermataceae [Limeaceae [[Lophocarpaceae [Barbeuiaceae [Aizoaceae

[Sarcobataceae Phytolaccaceae Nyctaginaceae]]]] [Molluginaceae [Halophytaceae

Montiaceae Basellaceae Didiereaceae [Talinaceae [Portulacaceae [Anacampseros etc +

Cactaceae]]]]]]]]]]]]]

ASTERIDS

Cornales

[Cornaceae + Nyssaceae] [Hydrangeaceae + Loasaceae] Hydrostachyaceae [Curtisiaceae

+ Grubbiaceae]

Ericales

[Balsaminaceae [Marcgraviaceae + Tetrameristaceae]] [[Polemoniaceae +

Fouquieraceae] Lecythidaceae [[Sladeniaceae + Pentaphylacaceae] [Sapotaceae

[Ebenaceae [Maesaceae [Theophrastaceae [Myrsinaceae + Primulaceae]]]]]

[Mitrastemonaceae Theaceae [Symplocaceae [Styracaceae + Diapensiaceae]]

[[Sarraceniaceae [Roridulaceae + Actinidiaceae] [Clethraceae [Cyrillaceae +

Ericaceae]]]]]]

LAMIIDASTERID I

Unplaced

Metteniusaceae Oncothecaceae Icacinaceae and some unplaced ex-Icacinaceae

Garryales

Garryaceae + Eucommiaceae

Unplaced

Boraginaceae Vahliaceae

Gentianales

Rubiaceae [Gentianaceae [Loganiaceae [Gelsemiaceae + Apocynaceae]]]

Lamiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

38

Plocospermataceae [Oleaceae [Tetrachondraceae [Calceolariaceae

[Gesneriaceae Peltanthera [Pedaliaceae Carlemanniaceae Martyniaceae Stilbaceae

Plantaginaceae Linderniaceae Bignoniaceae Verbenaceae Lamiaceae Paulowniaceae

Schlegeliaceae Thomandersiaceae Phrymaceae Rehmannia Acanthaceae

Scrophulariaceae Orobanchaceae Byblidaceae Lentibulariaceae]]]]]

Solanales

[Montiniaceae [Sphenocleaceae + Hydroleaceae]] [Convolvulaceae + Solanaceae]

CAMPANULIDASTERID II

Aquifoliales

[[Cardiopteridaceae + Stemonuraceae] [Aquifoliaceae [Helwingiaceae +

Phyllonomaceae]]

Asterales

[Rousseaceae + Campanulaceae] Pentaphragmataceae [[Alseuosmiaceae [Phellinaceae

+ Argophyllaceae]] Stylidiaceae [Menyanthaceae [Goodeniaceae [Calyceraceae +

Asteraceae]]]]

Escalloniales

Escalloniaceae

Bruniales

[Bruniaceae + Columelliaceae]

Apiales

Pennantiaceae [Torricelliaceae [Griseliniaceae [Pittosporaceae [Araliaceae

[Myodocarpaceae + Apiaceae]]]]]

Paracryphiales

Paracryphiaceae

Dipsacales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

39

Adoxaceae [Diervillaceae [Caprifoliaceae [Linnaeaceae [Morinaceae [Dipsacaceae +

Valerianaceae]]]]]

LINKS TO ORDERS AND FAMILIES

[Back to Top]

MAIN GROUPINGS

Asterids commelinids core eudicots asterid 1 asterid

2 eudicots gymnosperms Magnoliophyta monocots N-fixing claderosids seed plants

ALPHABETICAL LISTING OF ALL ORDINAL NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | L | M | N | O | P | Q | R | S | T | U | V | W | X | Z

Abietales Acanthales Acerales Acorales Actinidiales Actinostrobales Adoxales Aescul

ales Agavales Aizoales AkanialesAlismatales Alliales Alseuosmiales Alstroemeriales

Altingiales Amaranthales Amaryllidales Amborellales AmbrosialesAmmiales Amomal

es Ancistrocladales Anisophylleales Annonales Anthobolales Apiales Apocynales Apo

nogetonalesAquifoliales Arales Araliales Aralidiales Araucariales Arecales Aristolochi

ales Athrotaxidales Asarales AsclepiadalesAsparagales Asphodelales Asparagales Ast

eliales Atriplicales Aucubales Austrobaileyales Avenales

Balanitales Balanopales Balanophorales Balsaminales Barbeyales Barclayales Batales

Begoniales BerberidalesBerberidopsidales Betulales Biebersteiniales Bignoniales Bix

ales Boraginales Brassicales Brexiales Bromeliales BrunialesBrunoniales Burmannial

es Burserales Butomales Buxales Byblidales

Cactales Callitrichales Calycanthales Calycerales Campanulales Campynematales Can

ellales Cannales CapparalesCaprifoliales Cardiopteridales Carduales Caricales Carle

manniales Caryophyllales Cassiales Casuarinales CelastralesCentrolepidales Cephalot

ales Cephalotaxales Ceratophyllales Cercidiphyllales Chenopodiales Chironiales Chlor

anthalesChrysobalanales Cinchonales Circaeasterales Cistales Citrales Cocosales Col

chicales Columelliales CombretalesCommelinales Connarales Convolvulales Coriarial

es Cornales Corylales Corynocarpales Crassulales CrossosomatalesCucurbitales Cunn

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

40

inghamiales Cunoniales Cupressales Cycadales Cyclanthales Cymodoceales Cynarales

CynomorialesCyperales Cyrillales Cytinales

Daphnales Daphniphyllales Dasypogonales Datiscales Degeneriales Desfontainiales D

iapensiales Didymelales DillenialesDioales Dioncophyllales Dioscoreales Diospyrales

Dipentodontales Dipsacales Droserales

Ebenales Echiales Elaeagnales Elaeocarpales Elatinales Elodeales Emmotales Empetr

ales Ephedrales Ericales EriocaulalesErythropalaless Escalloniales Eucommiales Euph

orbiales Eupomatiales Eupteleales Euryalaless

Fabales Fagales Falcatifoliales Ficales Flacourtiales Flagellariales Fouquieriales Franc

oales Frangulales

Galiales Garryales Geissolomatales Gentianales Geraniales Gesneriales Ginkgoales G

laucidiales Globulariales GnetalesGoodeniales Greyiales Griseliniales Grossulariales

Grubbiales Gunnerales Gyrocarpales Gyrostemonales

Haemodorales Haloragales Hamamelidales Hanguanales Heisteriales Helleborales He

lwingiales Himantandrales HippuridalesHomaliales Hortensiales Huales Huerteales

Hydatellales Hydnorales Hydrangeales Hydrastidales HydrocharitalesHydropeltidales

Hydrostachyales Hypericales Hypoxidales

Icacinales Illiciales Iridales Irvingales Iteales Ixiales

Jasminales Juglandales Julianiales Juncaginales Juncales

Lacistematales Lactoridales Lamiales Lardizabalales Laurales Lecythidales Ledocarpal

es Leitneriales LentibularialesLigustrales Liliales Limnanthales Linales Liriales Loasal

es Lobeliales Loganiales Lonicerales Loranthales LowialesLythrales

Magnoliales Malpighiales Malvales Marathrales Marcgraviales Mayacales Medusagy

nales Medusandrales MelanthialesMelastomatales Meliales Melianthales Meliosmal

es Menispermales Menyanthales Metteniusales MitrastemonalesMiyoshiales Moni

miales Moringales Musales Myricales Myristicales Myrothamnales Myrsinales Myrta

les

Najadales Nandinales Narcissales Nartheciales Nelumbonales Nepenthales Neuradal

es Nitrariales Nolanales NothofagalesNyctaginales Nymphaeales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

41

Ochnales Oenotherales Olacales Oleales Onagrales Oncothecales Opuntiales Orchid

ales Oxalidales

Saxifragales Pandales Pandanales Papaverales Paracryphiales Parasitaxales Paridales

Parnassiales PassifloralesPenaeales Pennantiales Pentaphragmatales Peridiscales P

etiveriales Petrosaviales Phellinales Philydrales PhyllanthalesPhysenales Phytolaccal

es Picramniales Pinales Pinguiculales Piperales Pittosporales Plantaginales Platanale

sPlumbaginales Poales Podocarpales Podophyllales Podostemales Polemoniales Pol

ygalales Polygonales PontederialesPortulacales Posidoniales Potamogetonales Primu

lales Proteales

Quercales Quillajales Quintiniales

Rafflesiales Ranunculales Rapateales Resedales Restionales Rhabdodendrales Rhamn

ales Rhinanthales RhizophoralesRhodorales Rhoipteleales Roridulales Rosales Rouss

eales Rubiales Ruppiales Rutales

Sabiales Salicales Salvadorales Samolales Samydales Sanguisorbales Santalales Sapin

dales Sapotales SarracenialesSaxegotheales Saxifragales Scheuchzeriales Sciadopitya

les Scleranthales Scrophulariales Scyphostegiales SedalesSilenales Simmondsiales S

milacales Solanales Sphenocleales Sphenostemonales Stangeriales Stellariales Stemo

nalesStilbales Stylidiales Styracales Surianales

Taccales Tamales Tamaricales Taxales Taxodiales Tecophilaeales Terebinthales Tern

stroemiales Theales TheligonalesThymelaeales Tiliales Tofieldiales Torricelliales Tov

ariales Tribelales Trilliales Trimeniales Triuridales TrochodendralesTropaeolales Turn

erales Typhales

Ulmales Urticales

Vacciniales Vahliales Vallisneriales Velloziales Veratrales Verbenales Viburnales Vinc

ales Violales Viscales VitalesVochysiales

Welwitschiales Winterales

Xanthorrhoeales Ximeniales Xyridales

Zamiales Zingiberales Zosterales Zygophyllales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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ALPHABETICAL LISTING OF ALL FAMILY NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | K | L | M | N | O | P | Q | R | S | T | U | V | W | X | Y

|Z

Back to Home

In some cases clicking on a family synonym will send you directly to the subfamily in

which the genus of the basionym is to be found

Abietaceae Abolbodaceae Abrophyllaceae Acalyphaceae Acanthaceae [Acanthochla

mydaceae] Acarnaceaee AceraceaeAchariaceae Achatocarpaceae Achradaceae Achy

ranthaceae Acmopylaceae Acoraceae Acristaceae ActaeaceaeActinidiaceae Actinota

ceae Amentotaxaceae Adenogrammaceae Adoxaceae Aegialitidaceae Aegicerataceae

Aegilopaceae Aeginetiaceaee Aegiphilaceae Aesculaceae Aextoxicaceae Agapanthac

eae Agavaceae AgdestidaceaeAgialidaceae Agrimoniaceae Agrostidaceae Ailanthace

ae Aitoniaceae Aizoaceae Akaniaceae Alangiaceae AlchemillaceaeAldrovandaceae A

lismataceae Alliaceae Allioniaceae Allophylaceae Aloaceae Alopecuraceae Alpiniacea

e AlseuosmiaceaeAlsinaceae Alsodeiaceae Alstroemeriaceae Altingiaceae Alzateacea

e Amaranthaceae Amaryllidaceae Amborellaceae Ambrosiaceae Amentotaxaceae A

mmanniaceae Ammiaceae Amomaceae Ampelidopsaceae Amygdalaceae Amyridacea

eAnacardiaceae Anagallidaceae Anarthriaceae Anchusaceae Ancistrocladaceae Andr

omedaceae AndropogonaceaeAndrostachyaceae Androsynaceae Anemarrhenaceae

Anemonaceae Angelicaceae Anisophylleaceae AnnonaceaeAnomochloaceae Anopter

aceae Anrederaceae Anthemidaceae Anthericaceae Anthobolaceae Antidesmataceae

Antirrhinaceae Antoniaceae Aparinaceae Aphanopetalaceae Aphloiaceae Aphyllanth

aceae Apiaceae ApocynaceaeApodanthaceae Aponogetonaceae Aporusaceae Apose

ridaceae Apostasiaceae Aptandraceae AquifoliaceaeAquilariaceae Aquilegiaceae A

raceae Aragoaceae Araliaceae Aralidiaceaee Araucariaceae Arbutaceae Arceuthidace

aeArceuthobiaceae Arctostaphylaceae Arctotidaceae Ardisiaceae Arecaceae Argoph

yllaceae Arisaraceae AristolochiaceaeAristoteliaceae Arjonaceae Armeriaceae Artem

isiaceae Arthrotaxidaceae Artocarpaceae Arundinaceae ArundinellaceaeAsaraceae A

sclepiadaceae Ascyraceaee Asiraceae Asparagaceae Asperulaceae Asphodelaceae As

pidistraceaeeAsteliaceae Asteraceae Asteranthaceae Asteropeiaceae Astragalaceae

Astrocarpaceae AthanasiaceaeAtherospermataceae Atriplicaceae Atropaceae Aucuba

ceae Aurantiaceae Austrobaileyaceae Austrotaxaceae AvenaceaeAverrhoaceae Avet

raceae Avicenniaceae Azaleaceae Azimaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

43

Balanitaceae Balanopaceae Balanophoraceae Balsameaceae Balsaminaceae Bambusa

ceae Barbaceniaceae BarbeuiaceaeBarbeyaceae Barclayaceae Barringtoniaceae Base

llaceae Bataceae Baueraceae Bauhiniaceae Baxteriaceae BegoniaceaeBehniaceae B

elangeraceae Belloniaceae Bembiciaceae Berberidaceae Berberidopsidaceae Berryac

eaee BersamaceaeBertyaceae Berzeliaceae Besleriaceae Betaceae Betulaceae Biebe

rsteiniaceae Bifariaceae Bignoniaceae BischofiaceaeBixaceae Blakeaceae Blandfordia

ceae Blattiaceae Blepharocaryaceae Blitaceae Blyxaceae Boerlagellaceae Bolivaracea

eBombacaceae Bonnetiaceae Bontiaceae Boopidaceae Boraginaceae Borassaceae B

oroniaceae BoryaceaeBotryodendraceae Bougainvilleaceae Boweniaceae Brassicacea

e Brachycaulaceae Bretschneideraceae BrexiaceaeBromeliaceae Brownlowiaceae Br

unelliaceae Bruniaceae Brunoniaceae Brunsvigiaceae Bryoniaceae BuchneraceaeBuci

daceae Buddlejaceae Buglossaceae Bulbocodiaceae Bumeliaceae Bupleuraceae Burc

hardiaceae BurmanniaceaeBurseraceae Butneriaceae Butomaceae Buxaceae Byblida

ceae Byttneriaceae

Cabombaceae Cacaoaceae Cactaceae Caesalpiniaceae Caladiaceae Calamaceae Calce

olariaceae CalectasiaceaeCalendulaceae Callaceae Callicomaceae Calligonaceae Calli

traceae Callitrichaceae Calochortaceae CalophyllaceaeCalthaceae Calycanthaceae Ca

lyceraceae Cambogiaceae Camelliaceae Campanulaceae CampynemataceaeCanacom

yricaceae Canellaceae Cannabaceae Cannaceae Canopodaceae Canotiaceae Cansjera

ceae CapparaceaeCaprariaceae Caprifoliaceae Cardiopteridaceae Carduaceae Carica

ceae Carissaceae Carlemanniaceae CarpinaceaeCarpodetaceae Cartonemataceae Ca

ryocaraceae Caryophyllaceae Casearieaceae Cassiaceae CassipoureaceaeCassythacea

e Castaneaceae Castelaceae Casuarinaceae Catesbaeaceae Cathedraceae Cecropiace

ae CedraceaeCedrelaceae Celastraceae Celosiaceae Celtidaceae Centaureaceae Cen

trolepidaceae Centroplacaceae CepaceaeCephalanthaceae Cephalotaceae Cephalota

xaceae Cerastiaceae Ceratoniaceae Ceratophyllaceae CerberaceaeCercidiphyllaceae

Cercocarpaceae Cercodiaceae Cereaceae Cerinthaceae Ceroxylaceae Cestraceae Cev

alliaceaeChailletiaceae Chamaedoraceae Chamaemoraceae Chamelauciaceae Chaun

ochitonaceae Chelidoniaceae ChelonaceaeChenopodiaceae Chimonanthaceae Chingi

thamnaceae Chionographidaceae Chiranthodendraceae ChironiaceaeChlaenaceae Ch

loanthaceae Chloranthaceae Chloridaceae Chlorogalaceae Chrysobalanaceae Cicerac

eae CichoriaceaeCimicifugaceae Cinchoniaceae Circaeaceae Circaeasteraceae Cissac

eae Cistaceae Citraceae Clematidaceae CleomaceaeClethraceae Clusiaceae Cneorac

eae Cnestidaceae Cnicaceae Cobaeaceae Cochlospermaceae Cocosaceae Coffeaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

44

Colchicaceae Coleogynaceae Columelliaceae Combretaceae Commelinaceae Comocl

adiaceae Compositae CompsoaceaeConantheraceae Connaraceae Conostylidaceae C

onvallariaceae Convolvulaceae Cordiaceae CoreopsidaceaeCoriandraceae Coriariacea

e Coridaceae Corispermaceae Cornaceae Corokiaceae Coronillaceae Correaceae Cor

rigiolaceaeCorsiaceae Corydalaceae Corylaceae Corynocarpaceae Coryphaceae Cost

aceae Cotyledonaceae CoulaceaeCoutariaceae Coutoubiaceae Crassulaceae Crescen

tiaceae Cressaceae Crinaceae Crocaceae CroomiaceaeCrossosomataceae Crotonacea

e Cruciferae Cryptaceae Crypteroniaceae Cryptocorynaceae CryptomeriaceaeCtenol

ophonaceae Cucurbitaceae Cunninghamiaceae Cunoniaceae Cupressaceae Curcumac

eae Curtisiaceae CuscutaceaeCyananthaceae Cyanastraceae Cynanchaceae Cyanella

ceae Cycadaceae Cyclanthaceae CyclantheraceaeCyclocheilaceae Cydoniaceae Cymo

doceaceae Cynaraceae Cynocrambaceae Cynomoriaceae Cyperaceae CyphiaceaeCyp

hocarpaceae Cypripediaceae Cyrillaceae Cyrtandraceae Cyrtanthaceae Cytinaceae

Dacrycarpaceae Dactylanthaceae Damasoniaceae Daphnaceae Daphniphyllaceae Das

ypogonaceae DatiscaceaeDaturaceae Daucaceae Davidiaceae Davidsoniaceae Decais

neaceae Deeringiaceae Degeneriaceae DelphiniaceaeDendrophthoaceae Columelliac

eae Detariaceae Dialypetalanthaceae Dianellaceae Dianthaceae DiapensiaceaeDicha

petalaceae Dichondraceae Diclidantheraceae Dicrastylidiaceae Dictamnaceae Didiere

aceae DidymelaceaeDidymocarpaceae Diegodendraceae Diervillaceae Digitalidaceae

e Dilatridaceae Dilleniaceae Dioaceae DionaeaceaeDioncophyllaceae Dioscoreaceae

Diosmaceae Diospyraceae Dipentodontaceae Diphylleiaceae Diplolaenaceae Dipsaca

ceaeDipterocarpaceae Dirachmaceae Disanthaceae Diselmaceae Dodonaeaceae Do

mbeyaceae Donatiaceae DorsteniaceaeDortmannaceae Doryanthaceae Drabaceae D

racaenaceae Dracontiaceae Drimyidaceae Droseraceae DrosophyllaceaeDryadaceae

Duabangaceae Duckeodendraceae Dulongiaceae Durantaceae Durionaceae Dysphani

aceae

Ebenaceae Eccremidaceae Ecdeiocoleaceae Echiaceae Echinopaceae Ehretiaceae Ela

eagnaceae ElaeocarpaceaeElegiaceae Elatinaceae Ellisiaceae Ellisiophyllaceae Elodea

ceae Elytranthaceae Embeliaceae Emblingiaceae EmmotaceaeEmpetraceae Encepha

lartaceae Engelhardtiaceae Enhalaceae Epacridaceae Ephedraceae Ephemeranthacea

eEpilobiaceae Epimediaceae Eragrostidaceae Eremolepidaceae Eremosynaceae Erica

ceae Erinaceae EriocaulaceaeEriogonaceae Eriospermaceae Erodiaceae Erycibaceae

Eryngiaceae Erysimaceae Erythroniaceae ErythropalaceaeErythrospermaceae Erythro

xylaceae Escalloniaceae Eschscholziaceae Eucomidaceae Eucommiaceae Eucryphiace

aeEuonymaceae Eupatoriaceae Euphorbiaceae Euphrasiaceae Euphroniaceae Eupo

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

45

matiaceae Eupteleaceae EuryalaceaeEustrephaceae Euthemidaceae Eutocaceae Exb

ucklandiaceae Exocarpaceae

Fabaceae Fagaceae Falcatifoliaceae Ferulaceae Festucaceae Ficaceae Ficoidaceae Fit

zroyaceae FlacourtiaceaeFlagellariaceae Flindersiaceae Foetidiaceae Forestieraceae

Fothergillaceae Fouquieriaceae Fragariaceae FrancoaceaeFrangulaceae Frankeniacea

e Fraxinaceae Fraxinellaceae Freycinetiaceae Fritillariaceae Fuchsiaceae Fumariacea

eFunkiaceae

Gaiadendraceae Galacaceae Galanthaceae Galaxiaceae Galedupaceae Galeniaceae G

aliaceae Garciniaceae GardeniaceaeGardneriaceae Garryaceae Geissolomataceae Ge

itonoplesiaceae Gelsemiaceae Geniostomaceae GentianaceaeGeonomataceae Geosir

idaceae Geraniaceae Gerrardinaceae Gesneriaceae Gethyllidaceae Gilliesiaceae Gina

lloaceaeGinkgoaceae Gisekiaceae Gladiolaceae Glaucidiaceae Glechomaceae Glinace

ae Globulariaceae Gnaphaliaceae GnetaceaeGoetzeaceae Gomortegaceae Gomphiac

eae Gomphrenaceae Gonystylaceae Goodeniaceae Gordoniaceae GouaniaceaeGoup

iaceae Gramineae Gratiolaceae Grewiaceae Greyiaceae Grielaceae Griseliniaceae Gr

onoviaceae GrossulariaceaeGrubbiaceae Guaiacanaceae Guamatelaceae Guettardac

eae Gunneraceae Gustaviaceae Guttiferae GyrocarpaceaeGyrostemonaceae

Hachetteaceae Haemanthaceae Haemodoraceae Halesiaceae Hallieraceae Halocarpa

ceae Halophilaceae HalophytaceaeHaloragaceae Hamamelidaceae Hanguanaceae H

aptanthaceae Harmandaceae Hebenstretiaceae HectorellaceaeHederaceae Hedyosm

aceae Hedyotidaceae Hedysaraceae Heisteriaceae Heleniaceae Heliamphoraceae H

elianthaceaeHelianthemaceae Helichrysaceae Heliconiaceae Helicteraceae Heliotropi

aceae Helleboraceae Heloniadaceae HelosidaceaeHelwingiaceae Hemerocallidaceae

Hemimeridaceae Hemiphylacaceae Henriqueziaceae Henslowiaceae Hermanniaceae

Hernandiaceae Herniariaceae Herreriaceae Hesperocallidaceae Heterantheraceae He

teropyxidaceae HeterostylaceaeHibbertiaceae Hibiscaceae Hilleriaceae Himantandra

ceae Hippocastanaceae Hippocrateaceae HippomanaceaeHippophaeaceae Hippurida

ceae Hirtellaceae Holacanthaceae Homaliaceae Hopkinsiaceae Hoplestigmataceae H

ordeaceaeHornschuchiaceae Hortensiaceae Hortoniaceae Hostaceae Houstoniaceae

Huaceae Huerteaceae HugoniaceaeHumbertiaceae Humiriaceae Hyacinthaceae Hyd

atellaceae Hydnoraceae Hydrangeaceae Hydrastidaceae HydrillaceaeHydrocharitacea

e Hydrocotylaceae Hydrogetonaceae Hydroleaceae Hydropeltidaceae Hydrophylacac

eae HydrophyllaceaeHydrostachyaceae Hymenocardiaceae Hypecoaceae Hyperanth

eraceae Hypericaceae Hypopityaceae HypoxidaceaeHypseocharitaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

46

Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta

ceae ImperatoriaceaeInocarpaceae Inulaceae Ionidiaceae Iriarteaceae Iridaceae Irvi

ngiaceae Isnardiaceae Isophysidaceae IteaceaeStrasburgeriaceae Ixiaceae Ixioliriace

ae Ixonanthaceae

Jalapaceae Jamboliferaceae Japonoliriaceae Jasionaceae Jasminiaceae Johnsoniaceae

Joinvilleaceae JuglandaceaeJulianiaceae Juncaceae Juncaginaceae Juniperaceae Jus

siaeaceae Justiciaceae

Kadsuraceae Kaliphoraceae Kalmiaceae Kaniaceae Kiggelariaceae Kingdoniaceae Kin

giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae

Krameriaceae

Labiatae Lacandoniaceae Lachenaliaceae Lacistemataceae Lactoridaceae Lactucaceae

Lagerstroemiaceaee LamiaceaeLampsanaceae Lanariaceae Langsdorffiaceae Lantan

aceae Lapageriaceae Lardizabalaceae Lasiopetalaceae LauraceaeLawsoniaceae Laxm

anniaceae Lecythidaceae Ledaceae Ledocarpaceae Leeaceae Leguminosae Leitneriac

eae LemnaceaeLennoaceae Lentibulariaceae Lentiscaceae Leoniaceae Leonticaceae

Lepidobotryaceae LepidocarpaceaeLepidocaryaceae Lepidocerataceae Leptaulaceae

Leptospermaceae Lepturaceae Lepuropetalaceae LeuctenbergiaceaeLibocedraceae

Licaniaceae Ligustraceae Lilaeaceae Liliaceae Limeaceae Limnanthaceae Limnocharit

aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace

ae Linderniaceae Lindleyaceae Linnaeaceae LippayaceaeLiriaceae Liriodendraceae Li

ssocarpaceae Littorellaceae Loasaceae Lobeliaceaee Loganiaceae LomandraceaeLoni

ceraceae Lopeziaceae Lophiocarpaceae Lophiolaceae Lophiraceae Lophophytaceae L

ophopyxidaceae LoranthaceaeLotaceae Lowiaceae Lupulaceaee Luxemburgiaceae Lu

zuriagaceae Lychnidaceae Lyciaceae LyginiaceaeLygodisodeaceae Lysimachiaceae Ly

thraceae

Macarisiaceae Mackinlayaceae Madiaceae Maesaceae Magnoliaceae Malaceae Male

sherbiaceae MalortieaceaeMalpighiaceae Malvaceae Mangiaceae Manicariaceae Ma

paniaceae Marantaceae Marathraceae MarcgraviaceaeMartyniaceae Mastixiaceae

Matricariaceae Maundiaceae Mayacaceae Medeolaceae Medusagynaceae Peridiscac

eaeMelaleucaceae Melampyraceae Melanophyllaceae Melanthiaceae Melastomatac

eae Meliaceae Melianthaceae MelicaceaeMeliosmaceae Melittidaceae Melochiacea

e Memecylaceae Mendonciaceae Menispermaceae Menthaceae MenyanthaceaeMe

nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

47

Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

Nypaceae Nyssaceae

Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

ceae Papilionaceae Pappophoraceae PapyraceaeParacryphiaceae Parasitaxaceae Par

ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

48

hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

49

cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

50

ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 30: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

30

carried to clarify the distribution of morphological anatomical and chemical characters

for examples of what can to be done see the work of P K Endress and collaborators

(floral morphology and development) and S R Jensen and collaborators (iridoids) But

acquiring information about nodal anatomy cell and tissue distribution nodal anatomy

and the like is not difficult at all and should be generally encouraged

But all this is merely a necesary prelude to the understanding of evolution For this

one needs to know a lot more including dating the phylogeny understanding the

function(s) of characters factoring in relevant aspects of the palaeoenvironment in

which an apomorphy first appeared etc

SUMMARY OF THE SYSTEM

Below is a formalised summary of the relationships within orders of the families of seed

plants There are a few families that are not recognised even as options in APG II and

vice versa as well as a few extra orders The families to be recognised in the most

recent edition of Mabberleys The Plant Book (Mabberley 2008) are also largely

consistent with those below But all differences are trivial and will - I hope - eventually

disappear

Square brackets - [] - enclose clades the plus sign - + - designates sister taxa a

comma - - denotes part of a polytomy and quotation marks - - denotes a

paraphyletic group

SEED PLANTS

GYMNOSPERMS

Cycadales

Cycadaceae + Zamiaceae

Ginkgoales

Ginkgoaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

31

Pinales

Pinaceae [[Araucariaceae [Podocarpaceae + Phyllocladaceae]] [Sciadopityaceae

[Taxaceae + Cupressaceae]]]

Gnetales

Ephedraceae [Gnetaceae + Welwitschiaceae]

ANGIOSPERMSFLOWERING PLANTS

Amborellales

Amborellaceae

Nymphaeales

Hydatellaceae [Cabombaceae + Nymphaeaceae]

Austrobaileyales

Austrobaileyaceae [Schisandraceae + Trimeniaceae]

Chloranthales

Chloranthaceae

MAGNOLIIDS

Magnoliales

Myristicaceae [Magnoliaceae [[Himantandraceae + Degeneriaceae] [Eupomatiaceae +

Annonaceae]]]

Laurales

Calycanthaceae [[Siparunaceae [Gomortegaceae + Atherospermataceae]] [Monimiaceae

[Hernandiaceae + Lauraceae]]]

Canellales

Canellaceae + Winteraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

32

Piperales

[Hydnoraceae + Aristolochiaceae] [Piperaceae + Saururaceae]

MONOCOTS

Acorales

Acoraceae

Alismatales

Araceae [Tofieldiaceae [[Alismataceae [Hydrocharitaceae + Butomaceae]]

[Scheuchzeriaceae [Aponogetonaceae [Juncaginaceae [Maundiaceae [[Posidoniaceae

[Ruppiaceae + Cymodoceaceae]] [Zosteraceae + Potamogetonaceae]]]]]]]

Petrosaviales

Petrosaviaceae

Dioscoreales

Nartheciaceae [[Taccaceae + Thismiaceae] [Burmanniaceae + Dioscoreaceae]]

Pandanales

Velloziaceae Triuridaceae Stemonaceae [Pandanaceae + Cyclanthaceae]

Liliales

Corsiaceae [Campynemataceae [Petermanniaceae [Colchicaceae + Alstroemeriaceae]]

Melanthiaceae [[Philesiaceae + Rhipogonaceae] [Smilacaceae + Liliaceae]]]

Asparagales

Orchidaceae [[Boryaceae [Blandfordiaceae [Lanariaceae [Asteliaceae + Hypoxidaceae]]]]

[[Ixioliriaceae + Tecophilaeaceae] [Doryanthaceae [Iridaceae [Xeronemataceae

[Xanthorrhoeaceae [Alliaceae + Asparagaceae]]]]]]]

COMMELINIDS

Unplaced

Dasypogonaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

33

Arecales

Arecaceae

Poales

[Typhaceae + Bromeliaceae] [Rapateaceae [[Thurniaceae [Juncaceae + Cyperaceae]]

[[Mayacaceae [Eriocaulaceae + Xyridaceae]] [Flagellariaceae [Anarthriaceae

[Centrolepidaceae + Restionaceae]]] [Joinvilleaceae [Ecdeiocoleaceae + Poaceae]]]]]]]

Commelinales

[Commelinaceae + Hanguanaceae] [Philydraceae [Haemodoraceae + Pontederiaceae]]

Zingiberales

Musaceae [Strelitziaceae + Lowiaceae] Heliconiaceae [[Cannaceae + Marantaceae]

[Costaceae + Zingiberaceae]]

Ceratophyllales

Ceratophyllaceae

EUDICOTS

Ranunculales

Eupteleaceae [Papaveraceae [[[Lardizabalaceae + Circaeasteraceae] [Menispermaceae

[Berberidaceae + Ranunculaceae]]]]

Sabiales

Sabiaceae

Proteales

Nelumbonaceae [Platanaceae + Proteaceae]

Trochodendrales

Trochodendraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

34

Buxales

Haptanthaceae [Buxaceae + Didymelaceae]

CORE EUDICOTS

Gunnerales

Gunneraceae + Myrothamnaceae

Dilleniales

Dilleniaceae

Saxifragales

Peridiscaceae [[Paeoniaceae [Altingiaceae [Hamamelidaceae [Cercidiphyllaceae +

Daphniphyllaceae]]]] [[Crassulaceae [Aphanopetalaceae [Tetracarpaeaceae

[Penthoraceae + Haloragaceae]]]] [Iteaceae [Grossulariaceae + Saxifragaceae]]]]

Cynomoriaceae unplaced

Vitales

Vitaceae

ROSIDS

FABIDROSID I

Zygophyllales

Krameriaceae + Zygophyllaceae

Celastrales

Lepidobotryaceae + Celastraceae

Oxalidales

Huaceae [[Connaraceae + Oxalidaceae] [Cunoniaceae [Elaeocarpaceae [Brunelliaceae +

Cephalotaceae]]]]

Malpighiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

35

[Achariaceae [Goupiaceae [Violaceae + Passifloraceae] [Lacistemataceae Salicaceae]]

[Lophopyxidaceae Putranjivaceae] [Ctenolophonaceae [Erythroxylaceae +

Rhizophoraceae]] Linaceae Ixonanthaceae Humiriaceae Irvingiaceae

Centroplacaceae Pandaceae [[Bonnetiaceae + Clusiaceae] [Calophyllaceae

[Hypericaceae + Podostemaceae]]] [Malpighiaceae + Elatinaceae] Ochnaceae

[Peraceae [Rafflesiaceae + Euphorbiaceae]] [Phyllanthaceae + Picrodendraceae]

[Balanopaceae [[Trigoniaceae + Dichapetalaceae] [Chrysobalanaceae +

Euphroniaceae]]] Caryocaraceae

N-FIXING CLADE

Fabales

Quillajaceae [Fabaceae [Polygalaceae + Surianaceae]]

Rosales

Rosaceae [[Barbeyaceae [Elaeagnaceae Dirachmaceae Rhamnaceae]] [Ulmaceae

[Cannabaceae [Moraceae + Urticaceae]]]]

Cucurbitales

Anisophylleaceae [[Corynocarpaceae + Coriariaceae] [Cucurbitaceae [Tetramelaceae

[Datiscaceae + Begoniaceae]]]] Apodanthaceae

Fagales

Nothofagaceae [Fagaceae [[Myricaceae + Juglandaceae] [Casuarinaceae

[Ticodendraceae + Betulaceae]]]]

MALVIDROSID II

pgtGeraniales

Geraniaceae [[Melianthaceae + Francoaceae] [Vivianaceae + Ledocarpaceae]]

Myrtales

Combretaceae [[Onagraceae + Lythraceae] [[Vochysiaceae + Myrtaceae]

[Melastomataceae [Crypteroniaceae [Alzateaceae + Penaeaceae]]]]]

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

36

Crossosomatales

[Staphyleaceae [Guamatelaceae [Crossosomataceae + Stachyuraceae]]] [Aphloiaceae

[Geissolomataceae + Strasburgeriaceae]]

Picramniales

Picramniaceae

Sapindales

Biebersteiniaceae Nitrariaceae [Kirkiaceae [Anacardiaceae + Burseraceae]]

[Sapindaceae [Simaroubaceae Rutaceae Meliaceae]]

Huerteales

Gerradinaceae [Dipentodontaceae + Tapisciaceae]

Malvales

Neuradaceae [Thymelaeaceae [Sphaerosepalaceae Bixaceae [Cistaceae

[Sarcolaenaceae + Dipterocarpaceae]] [Cytinaceae + Muntingiaceae] Malvaceae]]

Brassicales

[Akaniaceae + Tropaeolaceae] [[Moringaceae + Caricaceae] [Setchellanthaceae

[Limnanthaceae [[Koeberliniaceae [Bataceae + Salvadoraceae]] [Emblingiaceae

[Pentadiplandraceae [Gyrostemonaceae Resedaceae Stixis and relatives] Tovariaceae

[Capparaceae [Cleomaceae + Brassicaceae]]]]]]]]

Berberidopsidales

Aextoxicaceae + Berberidopsidaceae

Santalales

Erythropalaceae Olacaceae [[Loranthaceae [Misodendraceae + Schoepfiaceae]]

[Opiliaceae + Santalaceae]] Balanophoraceae unplaced

Caryophyllales

[[Droseraceae [Nepenthaceae [Drosophyllaceae [Ancistrocladaceae +

Dioncophylleaceae]]]]] [[Frankeniaceae + Tamaricaceae] [Polygonaceae +

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

37

Plumbaginaceae]]] [Rhabdodendraceae [Simmondsiaceae [[Asteropeiaceae +

Physenaceae] [[Caryophyllaceae [Achatocarpaceae + Amaranthaceae]]

[Stegnospermataceae [Limeaceae [[Lophocarpaceae [Barbeuiaceae [Aizoaceae

[Sarcobataceae Phytolaccaceae Nyctaginaceae]]]] [Molluginaceae [Halophytaceae

Montiaceae Basellaceae Didiereaceae [Talinaceae [Portulacaceae [Anacampseros etc +

Cactaceae]]]]]]]]]]]]]

ASTERIDS

Cornales

[Cornaceae + Nyssaceae] [Hydrangeaceae + Loasaceae] Hydrostachyaceae [Curtisiaceae

+ Grubbiaceae]

Ericales

[Balsaminaceae [Marcgraviaceae + Tetrameristaceae]] [[Polemoniaceae +

Fouquieraceae] Lecythidaceae [[Sladeniaceae + Pentaphylacaceae] [Sapotaceae

[Ebenaceae [Maesaceae [Theophrastaceae [Myrsinaceae + Primulaceae]]]]]

[Mitrastemonaceae Theaceae [Symplocaceae [Styracaceae + Diapensiaceae]]

[[Sarraceniaceae [Roridulaceae + Actinidiaceae] [Clethraceae [Cyrillaceae +

Ericaceae]]]]]]

LAMIIDASTERID I

Unplaced

Metteniusaceae Oncothecaceae Icacinaceae and some unplaced ex-Icacinaceae

Garryales

Garryaceae + Eucommiaceae

Unplaced

Boraginaceae Vahliaceae

Gentianales

Rubiaceae [Gentianaceae [Loganiaceae [Gelsemiaceae + Apocynaceae]]]

Lamiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

38

Plocospermataceae [Oleaceae [Tetrachondraceae [Calceolariaceae

[Gesneriaceae Peltanthera [Pedaliaceae Carlemanniaceae Martyniaceae Stilbaceae

Plantaginaceae Linderniaceae Bignoniaceae Verbenaceae Lamiaceae Paulowniaceae

Schlegeliaceae Thomandersiaceae Phrymaceae Rehmannia Acanthaceae

Scrophulariaceae Orobanchaceae Byblidaceae Lentibulariaceae]]]]]

Solanales

[Montiniaceae [Sphenocleaceae + Hydroleaceae]] [Convolvulaceae + Solanaceae]

CAMPANULIDASTERID II

Aquifoliales

[[Cardiopteridaceae + Stemonuraceae] [Aquifoliaceae [Helwingiaceae +

Phyllonomaceae]]

Asterales

[Rousseaceae + Campanulaceae] Pentaphragmataceae [[Alseuosmiaceae [Phellinaceae

+ Argophyllaceae]] Stylidiaceae [Menyanthaceae [Goodeniaceae [Calyceraceae +

Asteraceae]]]]

Escalloniales

Escalloniaceae

Bruniales

[Bruniaceae + Columelliaceae]

Apiales

Pennantiaceae [Torricelliaceae [Griseliniaceae [Pittosporaceae [Araliaceae

[Myodocarpaceae + Apiaceae]]]]]

Paracryphiales

Paracryphiaceae

Dipsacales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

39

Adoxaceae [Diervillaceae [Caprifoliaceae [Linnaeaceae [Morinaceae [Dipsacaceae +

Valerianaceae]]]]]

LINKS TO ORDERS AND FAMILIES

[Back to Top]

MAIN GROUPINGS

Asterids commelinids core eudicots asterid 1 asterid

2 eudicots gymnosperms Magnoliophyta monocots N-fixing claderosids seed plants

ALPHABETICAL LISTING OF ALL ORDINAL NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | L | M | N | O | P | Q | R | S | T | U | V | W | X | Z

Abietales Acanthales Acerales Acorales Actinidiales Actinostrobales Adoxales Aescul

ales Agavales Aizoales AkanialesAlismatales Alliales Alseuosmiales Alstroemeriales

Altingiales Amaranthales Amaryllidales Amborellales AmbrosialesAmmiales Amomal

es Ancistrocladales Anisophylleales Annonales Anthobolales Apiales Apocynales Apo

nogetonalesAquifoliales Arales Araliales Aralidiales Araucariales Arecales Aristolochi

ales Athrotaxidales Asarales AsclepiadalesAsparagales Asphodelales Asparagales Ast

eliales Atriplicales Aucubales Austrobaileyales Avenales

Balanitales Balanopales Balanophorales Balsaminales Barbeyales Barclayales Batales

Begoniales BerberidalesBerberidopsidales Betulales Biebersteiniales Bignoniales Bix

ales Boraginales Brassicales Brexiales Bromeliales BrunialesBrunoniales Burmannial

es Burserales Butomales Buxales Byblidales

Cactales Callitrichales Calycanthales Calycerales Campanulales Campynematales Can

ellales Cannales CapparalesCaprifoliales Cardiopteridales Carduales Caricales Carle

manniales Caryophyllales Cassiales Casuarinales CelastralesCentrolepidales Cephalot

ales Cephalotaxales Ceratophyllales Cercidiphyllales Chenopodiales Chironiales Chlor

anthalesChrysobalanales Cinchonales Circaeasterales Cistales Citrales Cocosales Col

chicales Columelliales CombretalesCommelinales Connarales Convolvulales Coriarial

es Cornales Corylales Corynocarpales Crassulales CrossosomatalesCucurbitales Cunn

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

40

inghamiales Cunoniales Cupressales Cycadales Cyclanthales Cymodoceales Cynarales

CynomorialesCyperales Cyrillales Cytinales

Daphnales Daphniphyllales Dasypogonales Datiscales Degeneriales Desfontainiales D

iapensiales Didymelales DillenialesDioales Dioncophyllales Dioscoreales Diospyrales

Dipentodontales Dipsacales Droserales

Ebenales Echiales Elaeagnales Elaeocarpales Elatinales Elodeales Emmotales Empetr

ales Ephedrales Ericales EriocaulalesErythropalaless Escalloniales Eucommiales Euph

orbiales Eupomatiales Eupteleales Euryalaless

Fabales Fagales Falcatifoliales Ficales Flacourtiales Flagellariales Fouquieriales Franc

oales Frangulales

Galiales Garryales Geissolomatales Gentianales Geraniales Gesneriales Ginkgoales G

laucidiales Globulariales GnetalesGoodeniales Greyiales Griseliniales Grossulariales

Grubbiales Gunnerales Gyrocarpales Gyrostemonales

Haemodorales Haloragales Hamamelidales Hanguanales Heisteriales Helleborales He

lwingiales Himantandrales HippuridalesHomaliales Hortensiales Huales Huerteales

Hydatellales Hydnorales Hydrangeales Hydrastidales HydrocharitalesHydropeltidales

Hydrostachyales Hypericales Hypoxidales

Icacinales Illiciales Iridales Irvingales Iteales Ixiales

Jasminales Juglandales Julianiales Juncaginales Juncales

Lacistematales Lactoridales Lamiales Lardizabalales Laurales Lecythidales Ledocarpal

es Leitneriales LentibularialesLigustrales Liliales Limnanthales Linales Liriales Loasal

es Lobeliales Loganiales Lonicerales Loranthales LowialesLythrales

Magnoliales Malpighiales Malvales Marathrales Marcgraviales Mayacales Medusagy

nales Medusandrales MelanthialesMelastomatales Meliales Melianthales Meliosmal

es Menispermales Menyanthales Metteniusales MitrastemonalesMiyoshiales Moni

miales Moringales Musales Myricales Myristicales Myrothamnales Myrsinales Myrta

les

Najadales Nandinales Narcissales Nartheciales Nelumbonales Nepenthales Neuradal

es Nitrariales Nolanales NothofagalesNyctaginales Nymphaeales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

41

Ochnales Oenotherales Olacales Oleales Onagrales Oncothecales Opuntiales Orchid

ales Oxalidales

Saxifragales Pandales Pandanales Papaverales Paracryphiales Parasitaxales Paridales

Parnassiales PassifloralesPenaeales Pennantiales Pentaphragmatales Peridiscales P

etiveriales Petrosaviales Phellinales Philydrales PhyllanthalesPhysenales Phytolaccal

es Picramniales Pinales Pinguiculales Piperales Pittosporales Plantaginales Platanale

sPlumbaginales Poales Podocarpales Podophyllales Podostemales Polemoniales Pol

ygalales Polygonales PontederialesPortulacales Posidoniales Potamogetonales Primu

lales Proteales

Quercales Quillajales Quintiniales

Rafflesiales Ranunculales Rapateales Resedales Restionales Rhabdodendrales Rhamn

ales Rhinanthales RhizophoralesRhodorales Rhoipteleales Roridulales Rosales Rouss

eales Rubiales Ruppiales Rutales

Sabiales Salicales Salvadorales Samolales Samydales Sanguisorbales Santalales Sapin

dales Sapotales SarracenialesSaxegotheales Saxifragales Scheuchzeriales Sciadopitya

les Scleranthales Scrophulariales Scyphostegiales SedalesSilenales Simmondsiales S

milacales Solanales Sphenocleales Sphenostemonales Stangeriales Stellariales Stemo

nalesStilbales Stylidiales Styracales Surianales

Taccales Tamales Tamaricales Taxales Taxodiales Tecophilaeales Terebinthales Tern

stroemiales Theales TheligonalesThymelaeales Tiliales Tofieldiales Torricelliales Tov

ariales Tribelales Trilliales Trimeniales Triuridales TrochodendralesTropaeolales Turn

erales Typhales

Ulmales Urticales

Vacciniales Vahliales Vallisneriales Velloziales Veratrales Verbenales Viburnales Vinc

ales Violales Viscales VitalesVochysiales

Welwitschiales Winterales

Xanthorrhoeales Ximeniales Xyridales

Zamiales Zingiberales Zosterales Zygophyllales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

42

ALPHABETICAL LISTING OF ALL FAMILY NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | K | L | M | N | O | P | Q | R | S | T | U | V | W | X | Y

|Z

Back to Home

In some cases clicking on a family synonym will send you directly to the subfamily in

which the genus of the basionym is to be found

Abietaceae Abolbodaceae Abrophyllaceae Acalyphaceae Acanthaceae [Acanthochla

mydaceae] Acarnaceaee AceraceaeAchariaceae Achatocarpaceae Achradaceae Achy

ranthaceae Acmopylaceae Acoraceae Acristaceae ActaeaceaeActinidiaceae Actinota

ceae Amentotaxaceae Adenogrammaceae Adoxaceae Aegialitidaceae Aegicerataceae

Aegilopaceae Aeginetiaceaee Aegiphilaceae Aesculaceae Aextoxicaceae Agapanthac

eae Agavaceae AgdestidaceaeAgialidaceae Agrimoniaceae Agrostidaceae Ailanthace

ae Aitoniaceae Aizoaceae Akaniaceae Alangiaceae AlchemillaceaeAldrovandaceae A

lismataceae Alliaceae Allioniaceae Allophylaceae Aloaceae Alopecuraceae Alpiniacea

e AlseuosmiaceaeAlsinaceae Alsodeiaceae Alstroemeriaceae Altingiaceae Alzateacea

e Amaranthaceae Amaryllidaceae Amborellaceae Ambrosiaceae Amentotaxaceae A

mmanniaceae Ammiaceae Amomaceae Ampelidopsaceae Amygdalaceae Amyridacea

eAnacardiaceae Anagallidaceae Anarthriaceae Anchusaceae Ancistrocladaceae Andr

omedaceae AndropogonaceaeAndrostachyaceae Androsynaceae Anemarrhenaceae

Anemonaceae Angelicaceae Anisophylleaceae AnnonaceaeAnomochloaceae Anopter

aceae Anrederaceae Anthemidaceae Anthericaceae Anthobolaceae Antidesmataceae

Antirrhinaceae Antoniaceae Aparinaceae Aphanopetalaceae Aphloiaceae Aphyllanth

aceae Apiaceae ApocynaceaeApodanthaceae Aponogetonaceae Aporusaceae Apose

ridaceae Apostasiaceae Aptandraceae AquifoliaceaeAquilariaceae Aquilegiaceae A

raceae Aragoaceae Araliaceae Aralidiaceaee Araucariaceae Arbutaceae Arceuthidace

aeArceuthobiaceae Arctostaphylaceae Arctotidaceae Ardisiaceae Arecaceae Argoph

yllaceae Arisaraceae AristolochiaceaeAristoteliaceae Arjonaceae Armeriaceae Artem

isiaceae Arthrotaxidaceae Artocarpaceae Arundinaceae ArundinellaceaeAsaraceae A

sclepiadaceae Ascyraceaee Asiraceae Asparagaceae Asperulaceae Asphodelaceae As

pidistraceaeeAsteliaceae Asteraceae Asteranthaceae Asteropeiaceae Astragalaceae

Astrocarpaceae AthanasiaceaeAtherospermataceae Atriplicaceae Atropaceae Aucuba

ceae Aurantiaceae Austrobaileyaceae Austrotaxaceae AvenaceaeAverrhoaceae Avet

raceae Avicenniaceae Azaleaceae Azimaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

43

Balanitaceae Balanopaceae Balanophoraceae Balsameaceae Balsaminaceae Bambusa

ceae Barbaceniaceae BarbeuiaceaeBarbeyaceae Barclayaceae Barringtoniaceae Base

llaceae Bataceae Baueraceae Bauhiniaceae Baxteriaceae BegoniaceaeBehniaceae B

elangeraceae Belloniaceae Bembiciaceae Berberidaceae Berberidopsidaceae Berryac

eaee BersamaceaeBertyaceae Berzeliaceae Besleriaceae Betaceae Betulaceae Biebe

rsteiniaceae Bifariaceae Bignoniaceae BischofiaceaeBixaceae Blakeaceae Blandfordia

ceae Blattiaceae Blepharocaryaceae Blitaceae Blyxaceae Boerlagellaceae Bolivaracea

eBombacaceae Bonnetiaceae Bontiaceae Boopidaceae Boraginaceae Borassaceae B

oroniaceae BoryaceaeBotryodendraceae Bougainvilleaceae Boweniaceae Brassicacea

e Brachycaulaceae Bretschneideraceae BrexiaceaeBromeliaceae Brownlowiaceae Br

unelliaceae Bruniaceae Brunoniaceae Brunsvigiaceae Bryoniaceae BuchneraceaeBuci

daceae Buddlejaceae Buglossaceae Bulbocodiaceae Bumeliaceae Bupleuraceae Burc

hardiaceae BurmanniaceaeBurseraceae Butneriaceae Butomaceae Buxaceae Byblida

ceae Byttneriaceae

Cabombaceae Cacaoaceae Cactaceae Caesalpiniaceae Caladiaceae Calamaceae Calce

olariaceae CalectasiaceaeCalendulaceae Callaceae Callicomaceae Calligonaceae Calli

traceae Callitrichaceae Calochortaceae CalophyllaceaeCalthaceae Calycanthaceae Ca

lyceraceae Cambogiaceae Camelliaceae Campanulaceae CampynemataceaeCanacom

yricaceae Canellaceae Cannabaceae Cannaceae Canopodaceae Canotiaceae Cansjera

ceae CapparaceaeCaprariaceae Caprifoliaceae Cardiopteridaceae Carduaceae Carica

ceae Carissaceae Carlemanniaceae CarpinaceaeCarpodetaceae Cartonemataceae Ca

ryocaraceae Caryophyllaceae Casearieaceae Cassiaceae CassipoureaceaeCassythacea

e Castaneaceae Castelaceae Casuarinaceae Catesbaeaceae Cathedraceae Cecropiace

ae CedraceaeCedrelaceae Celastraceae Celosiaceae Celtidaceae Centaureaceae Cen

trolepidaceae Centroplacaceae CepaceaeCephalanthaceae Cephalotaceae Cephalota

xaceae Cerastiaceae Ceratoniaceae Ceratophyllaceae CerberaceaeCercidiphyllaceae

Cercocarpaceae Cercodiaceae Cereaceae Cerinthaceae Ceroxylaceae Cestraceae Cev

alliaceaeChailletiaceae Chamaedoraceae Chamaemoraceae Chamelauciaceae Chaun

ochitonaceae Chelidoniaceae ChelonaceaeChenopodiaceae Chimonanthaceae Chingi

thamnaceae Chionographidaceae Chiranthodendraceae ChironiaceaeChlaenaceae Ch

loanthaceae Chloranthaceae Chloridaceae Chlorogalaceae Chrysobalanaceae Cicerac

eae CichoriaceaeCimicifugaceae Cinchoniaceae Circaeaceae Circaeasteraceae Cissac

eae Cistaceae Citraceae Clematidaceae CleomaceaeClethraceae Clusiaceae Cneorac

eae Cnestidaceae Cnicaceae Cobaeaceae Cochlospermaceae Cocosaceae Coffeaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

44

Colchicaceae Coleogynaceae Columelliaceae Combretaceae Commelinaceae Comocl

adiaceae Compositae CompsoaceaeConantheraceae Connaraceae Conostylidaceae C

onvallariaceae Convolvulaceae Cordiaceae CoreopsidaceaeCoriandraceae Coriariacea

e Coridaceae Corispermaceae Cornaceae Corokiaceae Coronillaceae Correaceae Cor

rigiolaceaeCorsiaceae Corydalaceae Corylaceae Corynocarpaceae Coryphaceae Cost

aceae Cotyledonaceae CoulaceaeCoutariaceae Coutoubiaceae Crassulaceae Crescen

tiaceae Cressaceae Crinaceae Crocaceae CroomiaceaeCrossosomataceae Crotonacea

e Cruciferae Cryptaceae Crypteroniaceae Cryptocorynaceae CryptomeriaceaeCtenol

ophonaceae Cucurbitaceae Cunninghamiaceae Cunoniaceae Cupressaceae Curcumac

eae Curtisiaceae CuscutaceaeCyananthaceae Cyanastraceae Cynanchaceae Cyanella

ceae Cycadaceae Cyclanthaceae CyclantheraceaeCyclocheilaceae Cydoniaceae Cymo

doceaceae Cynaraceae Cynocrambaceae Cynomoriaceae Cyperaceae CyphiaceaeCyp

hocarpaceae Cypripediaceae Cyrillaceae Cyrtandraceae Cyrtanthaceae Cytinaceae

Dacrycarpaceae Dactylanthaceae Damasoniaceae Daphnaceae Daphniphyllaceae Das

ypogonaceae DatiscaceaeDaturaceae Daucaceae Davidiaceae Davidsoniaceae Decais

neaceae Deeringiaceae Degeneriaceae DelphiniaceaeDendrophthoaceae Columelliac

eae Detariaceae Dialypetalanthaceae Dianellaceae Dianthaceae DiapensiaceaeDicha

petalaceae Dichondraceae Diclidantheraceae Dicrastylidiaceae Dictamnaceae Didiere

aceae DidymelaceaeDidymocarpaceae Diegodendraceae Diervillaceae Digitalidaceae

e Dilatridaceae Dilleniaceae Dioaceae DionaeaceaeDioncophyllaceae Dioscoreaceae

Diosmaceae Diospyraceae Dipentodontaceae Diphylleiaceae Diplolaenaceae Dipsaca

ceaeDipterocarpaceae Dirachmaceae Disanthaceae Diselmaceae Dodonaeaceae Do

mbeyaceae Donatiaceae DorsteniaceaeDortmannaceae Doryanthaceae Drabaceae D

racaenaceae Dracontiaceae Drimyidaceae Droseraceae DrosophyllaceaeDryadaceae

Duabangaceae Duckeodendraceae Dulongiaceae Durantaceae Durionaceae Dysphani

aceae

Ebenaceae Eccremidaceae Ecdeiocoleaceae Echiaceae Echinopaceae Ehretiaceae Ela

eagnaceae ElaeocarpaceaeElegiaceae Elatinaceae Ellisiaceae Ellisiophyllaceae Elodea

ceae Elytranthaceae Embeliaceae Emblingiaceae EmmotaceaeEmpetraceae Encepha

lartaceae Engelhardtiaceae Enhalaceae Epacridaceae Ephedraceae Ephemeranthacea

eEpilobiaceae Epimediaceae Eragrostidaceae Eremolepidaceae Eremosynaceae Erica

ceae Erinaceae EriocaulaceaeEriogonaceae Eriospermaceae Erodiaceae Erycibaceae

Eryngiaceae Erysimaceae Erythroniaceae ErythropalaceaeErythrospermaceae Erythro

xylaceae Escalloniaceae Eschscholziaceae Eucomidaceae Eucommiaceae Eucryphiace

aeEuonymaceae Eupatoriaceae Euphorbiaceae Euphrasiaceae Euphroniaceae Eupo

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

45

matiaceae Eupteleaceae EuryalaceaeEustrephaceae Euthemidaceae Eutocaceae Exb

ucklandiaceae Exocarpaceae

Fabaceae Fagaceae Falcatifoliaceae Ferulaceae Festucaceae Ficaceae Ficoidaceae Fit

zroyaceae FlacourtiaceaeFlagellariaceae Flindersiaceae Foetidiaceae Forestieraceae

Fothergillaceae Fouquieriaceae Fragariaceae FrancoaceaeFrangulaceae Frankeniacea

e Fraxinaceae Fraxinellaceae Freycinetiaceae Fritillariaceae Fuchsiaceae Fumariacea

eFunkiaceae

Gaiadendraceae Galacaceae Galanthaceae Galaxiaceae Galedupaceae Galeniaceae G

aliaceae Garciniaceae GardeniaceaeGardneriaceae Garryaceae Geissolomataceae Ge

itonoplesiaceae Gelsemiaceae Geniostomaceae GentianaceaeGeonomataceae Geosir

idaceae Geraniaceae Gerrardinaceae Gesneriaceae Gethyllidaceae Gilliesiaceae Gina

lloaceaeGinkgoaceae Gisekiaceae Gladiolaceae Glaucidiaceae Glechomaceae Glinace

ae Globulariaceae Gnaphaliaceae GnetaceaeGoetzeaceae Gomortegaceae Gomphiac

eae Gomphrenaceae Gonystylaceae Goodeniaceae Gordoniaceae GouaniaceaeGoup

iaceae Gramineae Gratiolaceae Grewiaceae Greyiaceae Grielaceae Griseliniaceae Gr

onoviaceae GrossulariaceaeGrubbiaceae Guaiacanaceae Guamatelaceae Guettardac

eae Gunneraceae Gustaviaceae Guttiferae GyrocarpaceaeGyrostemonaceae

Hachetteaceae Haemanthaceae Haemodoraceae Halesiaceae Hallieraceae Halocarpa

ceae Halophilaceae HalophytaceaeHaloragaceae Hamamelidaceae Hanguanaceae H

aptanthaceae Harmandaceae Hebenstretiaceae HectorellaceaeHederaceae Hedyosm

aceae Hedyotidaceae Hedysaraceae Heisteriaceae Heleniaceae Heliamphoraceae H

elianthaceaeHelianthemaceae Helichrysaceae Heliconiaceae Helicteraceae Heliotropi

aceae Helleboraceae Heloniadaceae HelosidaceaeHelwingiaceae Hemerocallidaceae

Hemimeridaceae Hemiphylacaceae Henriqueziaceae Henslowiaceae Hermanniaceae

Hernandiaceae Herniariaceae Herreriaceae Hesperocallidaceae Heterantheraceae He

teropyxidaceae HeterostylaceaeHibbertiaceae Hibiscaceae Hilleriaceae Himantandra

ceae Hippocastanaceae Hippocrateaceae HippomanaceaeHippophaeaceae Hippurida

ceae Hirtellaceae Holacanthaceae Homaliaceae Hopkinsiaceae Hoplestigmataceae H

ordeaceaeHornschuchiaceae Hortensiaceae Hortoniaceae Hostaceae Houstoniaceae

Huaceae Huerteaceae HugoniaceaeHumbertiaceae Humiriaceae Hyacinthaceae Hyd

atellaceae Hydnoraceae Hydrangeaceae Hydrastidaceae HydrillaceaeHydrocharitacea

e Hydrocotylaceae Hydrogetonaceae Hydroleaceae Hydropeltidaceae Hydrophylacac

eae HydrophyllaceaeHydrostachyaceae Hymenocardiaceae Hypecoaceae Hyperanth

eraceae Hypericaceae Hypopityaceae HypoxidaceaeHypseocharitaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

46

Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta

ceae ImperatoriaceaeInocarpaceae Inulaceae Ionidiaceae Iriarteaceae Iridaceae Irvi

ngiaceae Isnardiaceae Isophysidaceae IteaceaeStrasburgeriaceae Ixiaceae Ixioliriace

ae Ixonanthaceae

Jalapaceae Jamboliferaceae Japonoliriaceae Jasionaceae Jasminiaceae Johnsoniaceae

Joinvilleaceae JuglandaceaeJulianiaceae Juncaceae Juncaginaceae Juniperaceae Jus

siaeaceae Justiciaceae

Kadsuraceae Kaliphoraceae Kalmiaceae Kaniaceae Kiggelariaceae Kingdoniaceae Kin

giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae

Krameriaceae

Labiatae Lacandoniaceae Lachenaliaceae Lacistemataceae Lactoridaceae Lactucaceae

Lagerstroemiaceaee LamiaceaeLampsanaceae Lanariaceae Langsdorffiaceae Lantan

aceae Lapageriaceae Lardizabalaceae Lasiopetalaceae LauraceaeLawsoniaceae Laxm

anniaceae Lecythidaceae Ledaceae Ledocarpaceae Leeaceae Leguminosae Leitneriac

eae LemnaceaeLennoaceae Lentibulariaceae Lentiscaceae Leoniaceae Leonticaceae

Lepidobotryaceae LepidocarpaceaeLepidocaryaceae Lepidocerataceae Leptaulaceae

Leptospermaceae Lepturaceae Lepuropetalaceae LeuctenbergiaceaeLibocedraceae

Licaniaceae Ligustraceae Lilaeaceae Liliaceae Limeaceae Limnanthaceae Limnocharit

aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace

ae Linderniaceae Lindleyaceae Linnaeaceae LippayaceaeLiriaceae Liriodendraceae Li

ssocarpaceae Littorellaceae Loasaceae Lobeliaceaee Loganiaceae LomandraceaeLoni

ceraceae Lopeziaceae Lophiocarpaceae Lophiolaceae Lophiraceae Lophophytaceae L

ophopyxidaceae LoranthaceaeLotaceae Lowiaceae Lupulaceaee Luxemburgiaceae Lu

zuriagaceae Lychnidaceae Lyciaceae LyginiaceaeLygodisodeaceae Lysimachiaceae Ly

thraceae

Macarisiaceae Mackinlayaceae Madiaceae Maesaceae Magnoliaceae Malaceae Male

sherbiaceae MalortieaceaeMalpighiaceae Malvaceae Mangiaceae Manicariaceae Ma

paniaceae Marantaceae Marathraceae MarcgraviaceaeMartyniaceae Mastixiaceae

Matricariaceae Maundiaceae Mayacaceae Medeolaceae Medusagynaceae Peridiscac

eaeMelaleucaceae Melampyraceae Melanophyllaceae Melanthiaceae Melastomatac

eae Meliaceae Melianthaceae MelicaceaeMeliosmaceae Melittidaceae Melochiacea

e Memecylaceae Mendonciaceae Menispermaceae Menthaceae MenyanthaceaeMe

nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

47

Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

Nypaceae Nyssaceae

Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

ceae Papilionaceae Pappophoraceae PapyraceaeParacryphiaceae Parasitaxaceae Par

ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

48

hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

49

cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

50

ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 31: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

31

Pinales

Pinaceae [[Araucariaceae [Podocarpaceae + Phyllocladaceae]] [Sciadopityaceae

[Taxaceae + Cupressaceae]]]

Gnetales

Ephedraceae [Gnetaceae + Welwitschiaceae]

ANGIOSPERMSFLOWERING PLANTS

Amborellales

Amborellaceae

Nymphaeales

Hydatellaceae [Cabombaceae + Nymphaeaceae]

Austrobaileyales

Austrobaileyaceae [Schisandraceae + Trimeniaceae]

Chloranthales

Chloranthaceae

MAGNOLIIDS

Magnoliales

Myristicaceae [Magnoliaceae [[Himantandraceae + Degeneriaceae] [Eupomatiaceae +

Annonaceae]]]

Laurales

Calycanthaceae [[Siparunaceae [Gomortegaceae + Atherospermataceae]] [Monimiaceae

[Hernandiaceae + Lauraceae]]]

Canellales

Canellaceae + Winteraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

32

Piperales

[Hydnoraceae + Aristolochiaceae] [Piperaceae + Saururaceae]

MONOCOTS

Acorales

Acoraceae

Alismatales

Araceae [Tofieldiaceae [[Alismataceae [Hydrocharitaceae + Butomaceae]]

[Scheuchzeriaceae [Aponogetonaceae [Juncaginaceae [Maundiaceae [[Posidoniaceae

[Ruppiaceae + Cymodoceaceae]] [Zosteraceae + Potamogetonaceae]]]]]]]

Petrosaviales

Petrosaviaceae

Dioscoreales

Nartheciaceae [[Taccaceae + Thismiaceae] [Burmanniaceae + Dioscoreaceae]]

Pandanales

Velloziaceae Triuridaceae Stemonaceae [Pandanaceae + Cyclanthaceae]

Liliales

Corsiaceae [Campynemataceae [Petermanniaceae [Colchicaceae + Alstroemeriaceae]]

Melanthiaceae [[Philesiaceae + Rhipogonaceae] [Smilacaceae + Liliaceae]]]

Asparagales

Orchidaceae [[Boryaceae [Blandfordiaceae [Lanariaceae [Asteliaceae + Hypoxidaceae]]]]

[[Ixioliriaceae + Tecophilaeaceae] [Doryanthaceae [Iridaceae [Xeronemataceae

[Xanthorrhoeaceae [Alliaceae + Asparagaceae]]]]]]]

COMMELINIDS

Unplaced

Dasypogonaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

33

Arecales

Arecaceae

Poales

[Typhaceae + Bromeliaceae] [Rapateaceae [[Thurniaceae [Juncaceae + Cyperaceae]]

[[Mayacaceae [Eriocaulaceae + Xyridaceae]] [Flagellariaceae [Anarthriaceae

[Centrolepidaceae + Restionaceae]]] [Joinvilleaceae [Ecdeiocoleaceae + Poaceae]]]]]]]

Commelinales

[Commelinaceae + Hanguanaceae] [Philydraceae [Haemodoraceae + Pontederiaceae]]

Zingiberales

Musaceae [Strelitziaceae + Lowiaceae] Heliconiaceae [[Cannaceae + Marantaceae]

[Costaceae + Zingiberaceae]]

Ceratophyllales

Ceratophyllaceae

EUDICOTS

Ranunculales

Eupteleaceae [Papaveraceae [[[Lardizabalaceae + Circaeasteraceae] [Menispermaceae

[Berberidaceae + Ranunculaceae]]]]

Sabiales

Sabiaceae

Proteales

Nelumbonaceae [Platanaceae + Proteaceae]

Trochodendrales

Trochodendraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

34

Buxales

Haptanthaceae [Buxaceae + Didymelaceae]

CORE EUDICOTS

Gunnerales

Gunneraceae + Myrothamnaceae

Dilleniales

Dilleniaceae

Saxifragales

Peridiscaceae [[Paeoniaceae [Altingiaceae [Hamamelidaceae [Cercidiphyllaceae +

Daphniphyllaceae]]]] [[Crassulaceae [Aphanopetalaceae [Tetracarpaeaceae

[Penthoraceae + Haloragaceae]]]] [Iteaceae [Grossulariaceae + Saxifragaceae]]]]

Cynomoriaceae unplaced

Vitales

Vitaceae

ROSIDS

FABIDROSID I

Zygophyllales

Krameriaceae + Zygophyllaceae

Celastrales

Lepidobotryaceae + Celastraceae

Oxalidales

Huaceae [[Connaraceae + Oxalidaceae] [Cunoniaceae [Elaeocarpaceae [Brunelliaceae +

Cephalotaceae]]]]

Malpighiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

35

[Achariaceae [Goupiaceae [Violaceae + Passifloraceae] [Lacistemataceae Salicaceae]]

[Lophopyxidaceae Putranjivaceae] [Ctenolophonaceae [Erythroxylaceae +

Rhizophoraceae]] Linaceae Ixonanthaceae Humiriaceae Irvingiaceae

Centroplacaceae Pandaceae [[Bonnetiaceae + Clusiaceae] [Calophyllaceae

[Hypericaceae + Podostemaceae]]] [Malpighiaceae + Elatinaceae] Ochnaceae

[Peraceae [Rafflesiaceae + Euphorbiaceae]] [Phyllanthaceae + Picrodendraceae]

[Balanopaceae [[Trigoniaceae + Dichapetalaceae] [Chrysobalanaceae +

Euphroniaceae]]] Caryocaraceae

N-FIXING CLADE

Fabales

Quillajaceae [Fabaceae [Polygalaceae + Surianaceae]]

Rosales

Rosaceae [[Barbeyaceae [Elaeagnaceae Dirachmaceae Rhamnaceae]] [Ulmaceae

[Cannabaceae [Moraceae + Urticaceae]]]]

Cucurbitales

Anisophylleaceae [[Corynocarpaceae + Coriariaceae] [Cucurbitaceae [Tetramelaceae

[Datiscaceae + Begoniaceae]]]] Apodanthaceae

Fagales

Nothofagaceae [Fagaceae [[Myricaceae + Juglandaceae] [Casuarinaceae

[Ticodendraceae + Betulaceae]]]]

MALVIDROSID II

pgtGeraniales

Geraniaceae [[Melianthaceae + Francoaceae] [Vivianaceae + Ledocarpaceae]]

Myrtales

Combretaceae [[Onagraceae + Lythraceae] [[Vochysiaceae + Myrtaceae]

[Melastomataceae [Crypteroniaceae [Alzateaceae + Penaeaceae]]]]]

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

36

Crossosomatales

[Staphyleaceae [Guamatelaceae [Crossosomataceae + Stachyuraceae]]] [Aphloiaceae

[Geissolomataceae + Strasburgeriaceae]]

Picramniales

Picramniaceae

Sapindales

Biebersteiniaceae Nitrariaceae [Kirkiaceae [Anacardiaceae + Burseraceae]]

[Sapindaceae [Simaroubaceae Rutaceae Meliaceae]]

Huerteales

Gerradinaceae [Dipentodontaceae + Tapisciaceae]

Malvales

Neuradaceae [Thymelaeaceae [Sphaerosepalaceae Bixaceae [Cistaceae

[Sarcolaenaceae + Dipterocarpaceae]] [Cytinaceae + Muntingiaceae] Malvaceae]]

Brassicales

[Akaniaceae + Tropaeolaceae] [[Moringaceae + Caricaceae] [Setchellanthaceae

[Limnanthaceae [[Koeberliniaceae [Bataceae + Salvadoraceae]] [Emblingiaceae

[Pentadiplandraceae [Gyrostemonaceae Resedaceae Stixis and relatives] Tovariaceae

[Capparaceae [Cleomaceae + Brassicaceae]]]]]]]]

Berberidopsidales

Aextoxicaceae + Berberidopsidaceae

Santalales

Erythropalaceae Olacaceae [[Loranthaceae [Misodendraceae + Schoepfiaceae]]

[Opiliaceae + Santalaceae]] Balanophoraceae unplaced

Caryophyllales

[[Droseraceae [Nepenthaceae [Drosophyllaceae [Ancistrocladaceae +

Dioncophylleaceae]]]]] [[Frankeniaceae + Tamaricaceae] [Polygonaceae +

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

37

Plumbaginaceae]]] [Rhabdodendraceae [Simmondsiaceae [[Asteropeiaceae +

Physenaceae] [[Caryophyllaceae [Achatocarpaceae + Amaranthaceae]]

[Stegnospermataceae [Limeaceae [[Lophocarpaceae [Barbeuiaceae [Aizoaceae

[Sarcobataceae Phytolaccaceae Nyctaginaceae]]]] [Molluginaceae [Halophytaceae

Montiaceae Basellaceae Didiereaceae [Talinaceae [Portulacaceae [Anacampseros etc +

Cactaceae]]]]]]]]]]]]]

ASTERIDS

Cornales

[Cornaceae + Nyssaceae] [Hydrangeaceae + Loasaceae] Hydrostachyaceae [Curtisiaceae

+ Grubbiaceae]

Ericales

[Balsaminaceae [Marcgraviaceae + Tetrameristaceae]] [[Polemoniaceae +

Fouquieraceae] Lecythidaceae [[Sladeniaceae + Pentaphylacaceae] [Sapotaceae

[Ebenaceae [Maesaceae [Theophrastaceae [Myrsinaceae + Primulaceae]]]]]

[Mitrastemonaceae Theaceae [Symplocaceae [Styracaceae + Diapensiaceae]]

[[Sarraceniaceae [Roridulaceae + Actinidiaceae] [Clethraceae [Cyrillaceae +

Ericaceae]]]]]]

LAMIIDASTERID I

Unplaced

Metteniusaceae Oncothecaceae Icacinaceae and some unplaced ex-Icacinaceae

Garryales

Garryaceae + Eucommiaceae

Unplaced

Boraginaceae Vahliaceae

Gentianales

Rubiaceae [Gentianaceae [Loganiaceae [Gelsemiaceae + Apocynaceae]]]

Lamiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

38

Plocospermataceae [Oleaceae [Tetrachondraceae [Calceolariaceae

[Gesneriaceae Peltanthera [Pedaliaceae Carlemanniaceae Martyniaceae Stilbaceae

Plantaginaceae Linderniaceae Bignoniaceae Verbenaceae Lamiaceae Paulowniaceae

Schlegeliaceae Thomandersiaceae Phrymaceae Rehmannia Acanthaceae

Scrophulariaceae Orobanchaceae Byblidaceae Lentibulariaceae]]]]]

Solanales

[Montiniaceae [Sphenocleaceae + Hydroleaceae]] [Convolvulaceae + Solanaceae]

CAMPANULIDASTERID II

Aquifoliales

[[Cardiopteridaceae + Stemonuraceae] [Aquifoliaceae [Helwingiaceae +

Phyllonomaceae]]

Asterales

[Rousseaceae + Campanulaceae] Pentaphragmataceae [[Alseuosmiaceae [Phellinaceae

+ Argophyllaceae]] Stylidiaceae [Menyanthaceae [Goodeniaceae [Calyceraceae +

Asteraceae]]]]

Escalloniales

Escalloniaceae

Bruniales

[Bruniaceae + Columelliaceae]

Apiales

Pennantiaceae [Torricelliaceae [Griseliniaceae [Pittosporaceae [Araliaceae

[Myodocarpaceae + Apiaceae]]]]]

Paracryphiales

Paracryphiaceae

Dipsacales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

39

Adoxaceae [Diervillaceae [Caprifoliaceae [Linnaeaceae [Morinaceae [Dipsacaceae +

Valerianaceae]]]]]

LINKS TO ORDERS AND FAMILIES

[Back to Top]

MAIN GROUPINGS

Asterids commelinids core eudicots asterid 1 asterid

2 eudicots gymnosperms Magnoliophyta monocots N-fixing claderosids seed plants

ALPHABETICAL LISTING OF ALL ORDINAL NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | L | M | N | O | P | Q | R | S | T | U | V | W | X | Z

Abietales Acanthales Acerales Acorales Actinidiales Actinostrobales Adoxales Aescul

ales Agavales Aizoales AkanialesAlismatales Alliales Alseuosmiales Alstroemeriales

Altingiales Amaranthales Amaryllidales Amborellales AmbrosialesAmmiales Amomal

es Ancistrocladales Anisophylleales Annonales Anthobolales Apiales Apocynales Apo

nogetonalesAquifoliales Arales Araliales Aralidiales Araucariales Arecales Aristolochi

ales Athrotaxidales Asarales AsclepiadalesAsparagales Asphodelales Asparagales Ast

eliales Atriplicales Aucubales Austrobaileyales Avenales

Balanitales Balanopales Balanophorales Balsaminales Barbeyales Barclayales Batales

Begoniales BerberidalesBerberidopsidales Betulales Biebersteiniales Bignoniales Bix

ales Boraginales Brassicales Brexiales Bromeliales BrunialesBrunoniales Burmannial

es Burserales Butomales Buxales Byblidales

Cactales Callitrichales Calycanthales Calycerales Campanulales Campynematales Can

ellales Cannales CapparalesCaprifoliales Cardiopteridales Carduales Caricales Carle

manniales Caryophyllales Cassiales Casuarinales CelastralesCentrolepidales Cephalot

ales Cephalotaxales Ceratophyllales Cercidiphyllales Chenopodiales Chironiales Chlor

anthalesChrysobalanales Cinchonales Circaeasterales Cistales Citrales Cocosales Col

chicales Columelliales CombretalesCommelinales Connarales Convolvulales Coriarial

es Cornales Corylales Corynocarpales Crassulales CrossosomatalesCucurbitales Cunn

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

40

inghamiales Cunoniales Cupressales Cycadales Cyclanthales Cymodoceales Cynarales

CynomorialesCyperales Cyrillales Cytinales

Daphnales Daphniphyllales Dasypogonales Datiscales Degeneriales Desfontainiales D

iapensiales Didymelales DillenialesDioales Dioncophyllales Dioscoreales Diospyrales

Dipentodontales Dipsacales Droserales

Ebenales Echiales Elaeagnales Elaeocarpales Elatinales Elodeales Emmotales Empetr

ales Ephedrales Ericales EriocaulalesErythropalaless Escalloniales Eucommiales Euph

orbiales Eupomatiales Eupteleales Euryalaless

Fabales Fagales Falcatifoliales Ficales Flacourtiales Flagellariales Fouquieriales Franc

oales Frangulales

Galiales Garryales Geissolomatales Gentianales Geraniales Gesneriales Ginkgoales G

laucidiales Globulariales GnetalesGoodeniales Greyiales Griseliniales Grossulariales

Grubbiales Gunnerales Gyrocarpales Gyrostemonales

Haemodorales Haloragales Hamamelidales Hanguanales Heisteriales Helleborales He

lwingiales Himantandrales HippuridalesHomaliales Hortensiales Huales Huerteales

Hydatellales Hydnorales Hydrangeales Hydrastidales HydrocharitalesHydropeltidales

Hydrostachyales Hypericales Hypoxidales

Icacinales Illiciales Iridales Irvingales Iteales Ixiales

Jasminales Juglandales Julianiales Juncaginales Juncales

Lacistematales Lactoridales Lamiales Lardizabalales Laurales Lecythidales Ledocarpal

es Leitneriales LentibularialesLigustrales Liliales Limnanthales Linales Liriales Loasal

es Lobeliales Loganiales Lonicerales Loranthales LowialesLythrales

Magnoliales Malpighiales Malvales Marathrales Marcgraviales Mayacales Medusagy

nales Medusandrales MelanthialesMelastomatales Meliales Melianthales Meliosmal

es Menispermales Menyanthales Metteniusales MitrastemonalesMiyoshiales Moni

miales Moringales Musales Myricales Myristicales Myrothamnales Myrsinales Myrta

les

Najadales Nandinales Narcissales Nartheciales Nelumbonales Nepenthales Neuradal

es Nitrariales Nolanales NothofagalesNyctaginales Nymphaeales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

41

Ochnales Oenotherales Olacales Oleales Onagrales Oncothecales Opuntiales Orchid

ales Oxalidales

Saxifragales Pandales Pandanales Papaverales Paracryphiales Parasitaxales Paridales

Parnassiales PassifloralesPenaeales Pennantiales Pentaphragmatales Peridiscales P

etiveriales Petrosaviales Phellinales Philydrales PhyllanthalesPhysenales Phytolaccal

es Picramniales Pinales Pinguiculales Piperales Pittosporales Plantaginales Platanale

sPlumbaginales Poales Podocarpales Podophyllales Podostemales Polemoniales Pol

ygalales Polygonales PontederialesPortulacales Posidoniales Potamogetonales Primu

lales Proteales

Quercales Quillajales Quintiniales

Rafflesiales Ranunculales Rapateales Resedales Restionales Rhabdodendrales Rhamn

ales Rhinanthales RhizophoralesRhodorales Rhoipteleales Roridulales Rosales Rouss

eales Rubiales Ruppiales Rutales

Sabiales Salicales Salvadorales Samolales Samydales Sanguisorbales Santalales Sapin

dales Sapotales SarracenialesSaxegotheales Saxifragales Scheuchzeriales Sciadopitya

les Scleranthales Scrophulariales Scyphostegiales SedalesSilenales Simmondsiales S

milacales Solanales Sphenocleales Sphenostemonales Stangeriales Stellariales Stemo

nalesStilbales Stylidiales Styracales Surianales

Taccales Tamales Tamaricales Taxales Taxodiales Tecophilaeales Terebinthales Tern

stroemiales Theales TheligonalesThymelaeales Tiliales Tofieldiales Torricelliales Tov

ariales Tribelales Trilliales Trimeniales Triuridales TrochodendralesTropaeolales Turn

erales Typhales

Ulmales Urticales

Vacciniales Vahliales Vallisneriales Velloziales Veratrales Verbenales Viburnales Vinc

ales Violales Viscales VitalesVochysiales

Welwitschiales Winterales

Xanthorrhoeales Ximeniales Xyridales

Zamiales Zingiberales Zosterales Zygophyllales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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ALPHABETICAL LISTING OF ALL FAMILY NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | K | L | M | N | O | P | Q | R | S | T | U | V | W | X | Y

|Z

Back to Home

In some cases clicking on a family synonym will send you directly to the subfamily in

which the genus of the basionym is to be found

Abietaceae Abolbodaceae Abrophyllaceae Acalyphaceae Acanthaceae [Acanthochla

mydaceae] Acarnaceaee AceraceaeAchariaceae Achatocarpaceae Achradaceae Achy

ranthaceae Acmopylaceae Acoraceae Acristaceae ActaeaceaeActinidiaceae Actinota

ceae Amentotaxaceae Adenogrammaceae Adoxaceae Aegialitidaceae Aegicerataceae

Aegilopaceae Aeginetiaceaee Aegiphilaceae Aesculaceae Aextoxicaceae Agapanthac

eae Agavaceae AgdestidaceaeAgialidaceae Agrimoniaceae Agrostidaceae Ailanthace

ae Aitoniaceae Aizoaceae Akaniaceae Alangiaceae AlchemillaceaeAldrovandaceae A

lismataceae Alliaceae Allioniaceae Allophylaceae Aloaceae Alopecuraceae Alpiniacea

e AlseuosmiaceaeAlsinaceae Alsodeiaceae Alstroemeriaceae Altingiaceae Alzateacea

e Amaranthaceae Amaryllidaceae Amborellaceae Ambrosiaceae Amentotaxaceae A

mmanniaceae Ammiaceae Amomaceae Ampelidopsaceae Amygdalaceae Amyridacea

eAnacardiaceae Anagallidaceae Anarthriaceae Anchusaceae Ancistrocladaceae Andr

omedaceae AndropogonaceaeAndrostachyaceae Androsynaceae Anemarrhenaceae

Anemonaceae Angelicaceae Anisophylleaceae AnnonaceaeAnomochloaceae Anopter

aceae Anrederaceae Anthemidaceae Anthericaceae Anthobolaceae Antidesmataceae

Antirrhinaceae Antoniaceae Aparinaceae Aphanopetalaceae Aphloiaceae Aphyllanth

aceae Apiaceae ApocynaceaeApodanthaceae Aponogetonaceae Aporusaceae Apose

ridaceae Apostasiaceae Aptandraceae AquifoliaceaeAquilariaceae Aquilegiaceae A

raceae Aragoaceae Araliaceae Aralidiaceaee Araucariaceae Arbutaceae Arceuthidace

aeArceuthobiaceae Arctostaphylaceae Arctotidaceae Ardisiaceae Arecaceae Argoph

yllaceae Arisaraceae AristolochiaceaeAristoteliaceae Arjonaceae Armeriaceae Artem

isiaceae Arthrotaxidaceae Artocarpaceae Arundinaceae ArundinellaceaeAsaraceae A

sclepiadaceae Ascyraceaee Asiraceae Asparagaceae Asperulaceae Asphodelaceae As

pidistraceaeeAsteliaceae Asteraceae Asteranthaceae Asteropeiaceae Astragalaceae

Astrocarpaceae AthanasiaceaeAtherospermataceae Atriplicaceae Atropaceae Aucuba

ceae Aurantiaceae Austrobaileyaceae Austrotaxaceae AvenaceaeAverrhoaceae Avet

raceae Avicenniaceae Azaleaceae Azimaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

43

Balanitaceae Balanopaceae Balanophoraceae Balsameaceae Balsaminaceae Bambusa

ceae Barbaceniaceae BarbeuiaceaeBarbeyaceae Barclayaceae Barringtoniaceae Base

llaceae Bataceae Baueraceae Bauhiniaceae Baxteriaceae BegoniaceaeBehniaceae B

elangeraceae Belloniaceae Bembiciaceae Berberidaceae Berberidopsidaceae Berryac

eaee BersamaceaeBertyaceae Berzeliaceae Besleriaceae Betaceae Betulaceae Biebe

rsteiniaceae Bifariaceae Bignoniaceae BischofiaceaeBixaceae Blakeaceae Blandfordia

ceae Blattiaceae Blepharocaryaceae Blitaceae Blyxaceae Boerlagellaceae Bolivaracea

eBombacaceae Bonnetiaceae Bontiaceae Boopidaceae Boraginaceae Borassaceae B

oroniaceae BoryaceaeBotryodendraceae Bougainvilleaceae Boweniaceae Brassicacea

e Brachycaulaceae Bretschneideraceae BrexiaceaeBromeliaceae Brownlowiaceae Br

unelliaceae Bruniaceae Brunoniaceae Brunsvigiaceae Bryoniaceae BuchneraceaeBuci

daceae Buddlejaceae Buglossaceae Bulbocodiaceae Bumeliaceae Bupleuraceae Burc

hardiaceae BurmanniaceaeBurseraceae Butneriaceae Butomaceae Buxaceae Byblida

ceae Byttneriaceae

Cabombaceae Cacaoaceae Cactaceae Caesalpiniaceae Caladiaceae Calamaceae Calce

olariaceae CalectasiaceaeCalendulaceae Callaceae Callicomaceae Calligonaceae Calli

traceae Callitrichaceae Calochortaceae CalophyllaceaeCalthaceae Calycanthaceae Ca

lyceraceae Cambogiaceae Camelliaceae Campanulaceae CampynemataceaeCanacom

yricaceae Canellaceae Cannabaceae Cannaceae Canopodaceae Canotiaceae Cansjera

ceae CapparaceaeCaprariaceae Caprifoliaceae Cardiopteridaceae Carduaceae Carica

ceae Carissaceae Carlemanniaceae CarpinaceaeCarpodetaceae Cartonemataceae Ca

ryocaraceae Caryophyllaceae Casearieaceae Cassiaceae CassipoureaceaeCassythacea

e Castaneaceae Castelaceae Casuarinaceae Catesbaeaceae Cathedraceae Cecropiace

ae CedraceaeCedrelaceae Celastraceae Celosiaceae Celtidaceae Centaureaceae Cen

trolepidaceae Centroplacaceae CepaceaeCephalanthaceae Cephalotaceae Cephalota

xaceae Cerastiaceae Ceratoniaceae Ceratophyllaceae CerberaceaeCercidiphyllaceae

Cercocarpaceae Cercodiaceae Cereaceae Cerinthaceae Ceroxylaceae Cestraceae Cev

alliaceaeChailletiaceae Chamaedoraceae Chamaemoraceae Chamelauciaceae Chaun

ochitonaceae Chelidoniaceae ChelonaceaeChenopodiaceae Chimonanthaceae Chingi

thamnaceae Chionographidaceae Chiranthodendraceae ChironiaceaeChlaenaceae Ch

loanthaceae Chloranthaceae Chloridaceae Chlorogalaceae Chrysobalanaceae Cicerac

eae CichoriaceaeCimicifugaceae Cinchoniaceae Circaeaceae Circaeasteraceae Cissac

eae Cistaceae Citraceae Clematidaceae CleomaceaeClethraceae Clusiaceae Cneorac

eae Cnestidaceae Cnicaceae Cobaeaceae Cochlospermaceae Cocosaceae Coffeaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Colchicaceae Coleogynaceae Columelliaceae Combretaceae Commelinaceae Comocl

adiaceae Compositae CompsoaceaeConantheraceae Connaraceae Conostylidaceae C

onvallariaceae Convolvulaceae Cordiaceae CoreopsidaceaeCoriandraceae Coriariacea

e Coridaceae Corispermaceae Cornaceae Corokiaceae Coronillaceae Correaceae Cor

rigiolaceaeCorsiaceae Corydalaceae Corylaceae Corynocarpaceae Coryphaceae Cost

aceae Cotyledonaceae CoulaceaeCoutariaceae Coutoubiaceae Crassulaceae Crescen

tiaceae Cressaceae Crinaceae Crocaceae CroomiaceaeCrossosomataceae Crotonacea

e Cruciferae Cryptaceae Crypteroniaceae Cryptocorynaceae CryptomeriaceaeCtenol

ophonaceae Cucurbitaceae Cunninghamiaceae Cunoniaceae Cupressaceae Curcumac

eae Curtisiaceae CuscutaceaeCyananthaceae Cyanastraceae Cynanchaceae Cyanella

ceae Cycadaceae Cyclanthaceae CyclantheraceaeCyclocheilaceae Cydoniaceae Cymo

doceaceae Cynaraceae Cynocrambaceae Cynomoriaceae Cyperaceae CyphiaceaeCyp

hocarpaceae Cypripediaceae Cyrillaceae Cyrtandraceae Cyrtanthaceae Cytinaceae

Dacrycarpaceae Dactylanthaceae Damasoniaceae Daphnaceae Daphniphyllaceae Das

ypogonaceae DatiscaceaeDaturaceae Daucaceae Davidiaceae Davidsoniaceae Decais

neaceae Deeringiaceae Degeneriaceae DelphiniaceaeDendrophthoaceae Columelliac

eae Detariaceae Dialypetalanthaceae Dianellaceae Dianthaceae DiapensiaceaeDicha

petalaceae Dichondraceae Diclidantheraceae Dicrastylidiaceae Dictamnaceae Didiere

aceae DidymelaceaeDidymocarpaceae Diegodendraceae Diervillaceae Digitalidaceae

e Dilatridaceae Dilleniaceae Dioaceae DionaeaceaeDioncophyllaceae Dioscoreaceae

Diosmaceae Diospyraceae Dipentodontaceae Diphylleiaceae Diplolaenaceae Dipsaca

ceaeDipterocarpaceae Dirachmaceae Disanthaceae Diselmaceae Dodonaeaceae Do

mbeyaceae Donatiaceae DorsteniaceaeDortmannaceae Doryanthaceae Drabaceae D

racaenaceae Dracontiaceae Drimyidaceae Droseraceae DrosophyllaceaeDryadaceae

Duabangaceae Duckeodendraceae Dulongiaceae Durantaceae Durionaceae Dysphani

aceae

Ebenaceae Eccremidaceae Ecdeiocoleaceae Echiaceae Echinopaceae Ehretiaceae Ela

eagnaceae ElaeocarpaceaeElegiaceae Elatinaceae Ellisiaceae Ellisiophyllaceae Elodea

ceae Elytranthaceae Embeliaceae Emblingiaceae EmmotaceaeEmpetraceae Encepha

lartaceae Engelhardtiaceae Enhalaceae Epacridaceae Ephedraceae Ephemeranthacea

eEpilobiaceae Epimediaceae Eragrostidaceae Eremolepidaceae Eremosynaceae Erica

ceae Erinaceae EriocaulaceaeEriogonaceae Eriospermaceae Erodiaceae Erycibaceae

Eryngiaceae Erysimaceae Erythroniaceae ErythropalaceaeErythrospermaceae Erythro

xylaceae Escalloniaceae Eschscholziaceae Eucomidaceae Eucommiaceae Eucryphiace

aeEuonymaceae Eupatoriaceae Euphorbiaceae Euphrasiaceae Euphroniaceae Eupo

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

45

matiaceae Eupteleaceae EuryalaceaeEustrephaceae Euthemidaceae Eutocaceae Exb

ucklandiaceae Exocarpaceae

Fabaceae Fagaceae Falcatifoliaceae Ferulaceae Festucaceae Ficaceae Ficoidaceae Fit

zroyaceae FlacourtiaceaeFlagellariaceae Flindersiaceae Foetidiaceae Forestieraceae

Fothergillaceae Fouquieriaceae Fragariaceae FrancoaceaeFrangulaceae Frankeniacea

e Fraxinaceae Fraxinellaceae Freycinetiaceae Fritillariaceae Fuchsiaceae Fumariacea

eFunkiaceae

Gaiadendraceae Galacaceae Galanthaceae Galaxiaceae Galedupaceae Galeniaceae G

aliaceae Garciniaceae GardeniaceaeGardneriaceae Garryaceae Geissolomataceae Ge

itonoplesiaceae Gelsemiaceae Geniostomaceae GentianaceaeGeonomataceae Geosir

idaceae Geraniaceae Gerrardinaceae Gesneriaceae Gethyllidaceae Gilliesiaceae Gina

lloaceaeGinkgoaceae Gisekiaceae Gladiolaceae Glaucidiaceae Glechomaceae Glinace

ae Globulariaceae Gnaphaliaceae GnetaceaeGoetzeaceae Gomortegaceae Gomphiac

eae Gomphrenaceae Gonystylaceae Goodeniaceae Gordoniaceae GouaniaceaeGoup

iaceae Gramineae Gratiolaceae Grewiaceae Greyiaceae Grielaceae Griseliniaceae Gr

onoviaceae GrossulariaceaeGrubbiaceae Guaiacanaceae Guamatelaceae Guettardac

eae Gunneraceae Gustaviaceae Guttiferae GyrocarpaceaeGyrostemonaceae

Hachetteaceae Haemanthaceae Haemodoraceae Halesiaceae Hallieraceae Halocarpa

ceae Halophilaceae HalophytaceaeHaloragaceae Hamamelidaceae Hanguanaceae H

aptanthaceae Harmandaceae Hebenstretiaceae HectorellaceaeHederaceae Hedyosm

aceae Hedyotidaceae Hedysaraceae Heisteriaceae Heleniaceae Heliamphoraceae H

elianthaceaeHelianthemaceae Helichrysaceae Heliconiaceae Helicteraceae Heliotropi

aceae Helleboraceae Heloniadaceae HelosidaceaeHelwingiaceae Hemerocallidaceae

Hemimeridaceae Hemiphylacaceae Henriqueziaceae Henslowiaceae Hermanniaceae

Hernandiaceae Herniariaceae Herreriaceae Hesperocallidaceae Heterantheraceae He

teropyxidaceae HeterostylaceaeHibbertiaceae Hibiscaceae Hilleriaceae Himantandra

ceae Hippocastanaceae Hippocrateaceae HippomanaceaeHippophaeaceae Hippurida

ceae Hirtellaceae Holacanthaceae Homaliaceae Hopkinsiaceae Hoplestigmataceae H

ordeaceaeHornschuchiaceae Hortensiaceae Hortoniaceae Hostaceae Houstoniaceae

Huaceae Huerteaceae HugoniaceaeHumbertiaceae Humiriaceae Hyacinthaceae Hyd

atellaceae Hydnoraceae Hydrangeaceae Hydrastidaceae HydrillaceaeHydrocharitacea

e Hydrocotylaceae Hydrogetonaceae Hydroleaceae Hydropeltidaceae Hydrophylacac

eae HydrophyllaceaeHydrostachyaceae Hymenocardiaceae Hypecoaceae Hyperanth

eraceae Hypericaceae Hypopityaceae HypoxidaceaeHypseocharitaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta

ceae ImperatoriaceaeInocarpaceae Inulaceae Ionidiaceae Iriarteaceae Iridaceae Irvi

ngiaceae Isnardiaceae Isophysidaceae IteaceaeStrasburgeriaceae Ixiaceae Ixioliriace

ae Ixonanthaceae

Jalapaceae Jamboliferaceae Japonoliriaceae Jasionaceae Jasminiaceae Johnsoniaceae

Joinvilleaceae JuglandaceaeJulianiaceae Juncaceae Juncaginaceae Juniperaceae Jus

siaeaceae Justiciaceae

Kadsuraceae Kaliphoraceae Kalmiaceae Kaniaceae Kiggelariaceae Kingdoniaceae Kin

giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae

Krameriaceae

Labiatae Lacandoniaceae Lachenaliaceae Lacistemataceae Lactoridaceae Lactucaceae

Lagerstroemiaceaee LamiaceaeLampsanaceae Lanariaceae Langsdorffiaceae Lantan

aceae Lapageriaceae Lardizabalaceae Lasiopetalaceae LauraceaeLawsoniaceae Laxm

anniaceae Lecythidaceae Ledaceae Ledocarpaceae Leeaceae Leguminosae Leitneriac

eae LemnaceaeLennoaceae Lentibulariaceae Lentiscaceae Leoniaceae Leonticaceae

Lepidobotryaceae LepidocarpaceaeLepidocaryaceae Lepidocerataceae Leptaulaceae

Leptospermaceae Lepturaceae Lepuropetalaceae LeuctenbergiaceaeLibocedraceae

Licaniaceae Ligustraceae Lilaeaceae Liliaceae Limeaceae Limnanthaceae Limnocharit

aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace

ae Linderniaceae Lindleyaceae Linnaeaceae LippayaceaeLiriaceae Liriodendraceae Li

ssocarpaceae Littorellaceae Loasaceae Lobeliaceaee Loganiaceae LomandraceaeLoni

ceraceae Lopeziaceae Lophiocarpaceae Lophiolaceae Lophiraceae Lophophytaceae L

ophopyxidaceae LoranthaceaeLotaceae Lowiaceae Lupulaceaee Luxemburgiaceae Lu

zuriagaceae Lychnidaceae Lyciaceae LyginiaceaeLygodisodeaceae Lysimachiaceae Ly

thraceae

Macarisiaceae Mackinlayaceae Madiaceae Maesaceae Magnoliaceae Malaceae Male

sherbiaceae MalortieaceaeMalpighiaceae Malvaceae Mangiaceae Manicariaceae Ma

paniaceae Marantaceae Marathraceae MarcgraviaceaeMartyniaceae Mastixiaceae

Matricariaceae Maundiaceae Mayacaceae Medeolaceae Medusagynaceae Peridiscac

eaeMelaleucaceae Melampyraceae Melanophyllaceae Melanthiaceae Melastomatac

eae Meliaceae Melianthaceae MelicaceaeMeliosmaceae Melittidaceae Melochiacea

e Memecylaceae Mendonciaceae Menispermaceae Menthaceae MenyanthaceaeMe

nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

47

Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

Nypaceae Nyssaceae

Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

ceae Papilionaceae Pappophoraceae PapyraceaeParacryphiaceae Parasitaxaceae Par

ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

48

hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

49

cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

50

ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 32: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

32

Piperales

[Hydnoraceae + Aristolochiaceae] [Piperaceae + Saururaceae]

MONOCOTS

Acorales

Acoraceae

Alismatales

Araceae [Tofieldiaceae [[Alismataceae [Hydrocharitaceae + Butomaceae]]

[Scheuchzeriaceae [Aponogetonaceae [Juncaginaceae [Maundiaceae [[Posidoniaceae

[Ruppiaceae + Cymodoceaceae]] [Zosteraceae + Potamogetonaceae]]]]]]]

Petrosaviales

Petrosaviaceae

Dioscoreales

Nartheciaceae [[Taccaceae + Thismiaceae] [Burmanniaceae + Dioscoreaceae]]

Pandanales

Velloziaceae Triuridaceae Stemonaceae [Pandanaceae + Cyclanthaceae]

Liliales

Corsiaceae [Campynemataceae [Petermanniaceae [Colchicaceae + Alstroemeriaceae]]

Melanthiaceae [[Philesiaceae + Rhipogonaceae] [Smilacaceae + Liliaceae]]]

Asparagales

Orchidaceae [[Boryaceae [Blandfordiaceae [Lanariaceae [Asteliaceae + Hypoxidaceae]]]]

[[Ixioliriaceae + Tecophilaeaceae] [Doryanthaceae [Iridaceae [Xeronemataceae

[Xanthorrhoeaceae [Alliaceae + Asparagaceae]]]]]]]

COMMELINIDS

Unplaced

Dasypogonaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

33

Arecales

Arecaceae

Poales

[Typhaceae + Bromeliaceae] [Rapateaceae [[Thurniaceae [Juncaceae + Cyperaceae]]

[[Mayacaceae [Eriocaulaceae + Xyridaceae]] [Flagellariaceae [Anarthriaceae

[Centrolepidaceae + Restionaceae]]] [Joinvilleaceae [Ecdeiocoleaceae + Poaceae]]]]]]]

Commelinales

[Commelinaceae + Hanguanaceae] [Philydraceae [Haemodoraceae + Pontederiaceae]]

Zingiberales

Musaceae [Strelitziaceae + Lowiaceae] Heliconiaceae [[Cannaceae + Marantaceae]

[Costaceae + Zingiberaceae]]

Ceratophyllales

Ceratophyllaceae

EUDICOTS

Ranunculales

Eupteleaceae [Papaveraceae [[[Lardizabalaceae + Circaeasteraceae] [Menispermaceae

[Berberidaceae + Ranunculaceae]]]]

Sabiales

Sabiaceae

Proteales

Nelumbonaceae [Platanaceae + Proteaceae]

Trochodendrales

Trochodendraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

34

Buxales

Haptanthaceae [Buxaceae + Didymelaceae]

CORE EUDICOTS

Gunnerales

Gunneraceae + Myrothamnaceae

Dilleniales

Dilleniaceae

Saxifragales

Peridiscaceae [[Paeoniaceae [Altingiaceae [Hamamelidaceae [Cercidiphyllaceae +

Daphniphyllaceae]]]] [[Crassulaceae [Aphanopetalaceae [Tetracarpaeaceae

[Penthoraceae + Haloragaceae]]]] [Iteaceae [Grossulariaceae + Saxifragaceae]]]]

Cynomoriaceae unplaced

Vitales

Vitaceae

ROSIDS

FABIDROSID I

Zygophyllales

Krameriaceae + Zygophyllaceae

Celastrales

Lepidobotryaceae + Celastraceae

Oxalidales

Huaceae [[Connaraceae + Oxalidaceae] [Cunoniaceae [Elaeocarpaceae [Brunelliaceae +

Cephalotaceae]]]]

Malpighiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

35

[Achariaceae [Goupiaceae [Violaceae + Passifloraceae] [Lacistemataceae Salicaceae]]

[Lophopyxidaceae Putranjivaceae] [Ctenolophonaceae [Erythroxylaceae +

Rhizophoraceae]] Linaceae Ixonanthaceae Humiriaceae Irvingiaceae

Centroplacaceae Pandaceae [[Bonnetiaceae + Clusiaceae] [Calophyllaceae

[Hypericaceae + Podostemaceae]]] [Malpighiaceae + Elatinaceae] Ochnaceae

[Peraceae [Rafflesiaceae + Euphorbiaceae]] [Phyllanthaceae + Picrodendraceae]

[Balanopaceae [[Trigoniaceae + Dichapetalaceae] [Chrysobalanaceae +

Euphroniaceae]]] Caryocaraceae

N-FIXING CLADE

Fabales

Quillajaceae [Fabaceae [Polygalaceae + Surianaceae]]

Rosales

Rosaceae [[Barbeyaceae [Elaeagnaceae Dirachmaceae Rhamnaceae]] [Ulmaceae

[Cannabaceae [Moraceae + Urticaceae]]]]

Cucurbitales

Anisophylleaceae [[Corynocarpaceae + Coriariaceae] [Cucurbitaceae [Tetramelaceae

[Datiscaceae + Begoniaceae]]]] Apodanthaceae

Fagales

Nothofagaceae [Fagaceae [[Myricaceae + Juglandaceae] [Casuarinaceae

[Ticodendraceae + Betulaceae]]]]

MALVIDROSID II

pgtGeraniales

Geraniaceae [[Melianthaceae + Francoaceae] [Vivianaceae + Ledocarpaceae]]

Myrtales

Combretaceae [[Onagraceae + Lythraceae] [[Vochysiaceae + Myrtaceae]

[Melastomataceae [Crypteroniaceae [Alzateaceae + Penaeaceae]]]]]

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

36

Crossosomatales

[Staphyleaceae [Guamatelaceae [Crossosomataceae + Stachyuraceae]]] [Aphloiaceae

[Geissolomataceae + Strasburgeriaceae]]

Picramniales

Picramniaceae

Sapindales

Biebersteiniaceae Nitrariaceae [Kirkiaceae [Anacardiaceae + Burseraceae]]

[Sapindaceae [Simaroubaceae Rutaceae Meliaceae]]

Huerteales

Gerradinaceae [Dipentodontaceae + Tapisciaceae]

Malvales

Neuradaceae [Thymelaeaceae [Sphaerosepalaceae Bixaceae [Cistaceae

[Sarcolaenaceae + Dipterocarpaceae]] [Cytinaceae + Muntingiaceae] Malvaceae]]

Brassicales

[Akaniaceae + Tropaeolaceae] [[Moringaceae + Caricaceae] [Setchellanthaceae

[Limnanthaceae [[Koeberliniaceae [Bataceae + Salvadoraceae]] [Emblingiaceae

[Pentadiplandraceae [Gyrostemonaceae Resedaceae Stixis and relatives] Tovariaceae

[Capparaceae [Cleomaceae + Brassicaceae]]]]]]]]

Berberidopsidales

Aextoxicaceae + Berberidopsidaceae

Santalales

Erythropalaceae Olacaceae [[Loranthaceae [Misodendraceae + Schoepfiaceae]]

[Opiliaceae + Santalaceae]] Balanophoraceae unplaced

Caryophyllales

[[Droseraceae [Nepenthaceae [Drosophyllaceae [Ancistrocladaceae +

Dioncophylleaceae]]]]] [[Frankeniaceae + Tamaricaceae] [Polygonaceae +

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

37

Plumbaginaceae]]] [Rhabdodendraceae [Simmondsiaceae [[Asteropeiaceae +

Physenaceae] [[Caryophyllaceae [Achatocarpaceae + Amaranthaceae]]

[Stegnospermataceae [Limeaceae [[Lophocarpaceae [Barbeuiaceae [Aizoaceae

[Sarcobataceae Phytolaccaceae Nyctaginaceae]]]] [Molluginaceae [Halophytaceae

Montiaceae Basellaceae Didiereaceae [Talinaceae [Portulacaceae [Anacampseros etc +

Cactaceae]]]]]]]]]]]]]

ASTERIDS

Cornales

[Cornaceae + Nyssaceae] [Hydrangeaceae + Loasaceae] Hydrostachyaceae [Curtisiaceae

+ Grubbiaceae]

Ericales

[Balsaminaceae [Marcgraviaceae + Tetrameristaceae]] [[Polemoniaceae +

Fouquieraceae] Lecythidaceae [[Sladeniaceae + Pentaphylacaceae] [Sapotaceae

[Ebenaceae [Maesaceae [Theophrastaceae [Myrsinaceae + Primulaceae]]]]]

[Mitrastemonaceae Theaceae [Symplocaceae [Styracaceae + Diapensiaceae]]

[[Sarraceniaceae [Roridulaceae + Actinidiaceae] [Clethraceae [Cyrillaceae +

Ericaceae]]]]]]

LAMIIDASTERID I

Unplaced

Metteniusaceae Oncothecaceae Icacinaceae and some unplaced ex-Icacinaceae

Garryales

Garryaceae + Eucommiaceae

Unplaced

Boraginaceae Vahliaceae

Gentianales

Rubiaceae [Gentianaceae [Loganiaceae [Gelsemiaceae + Apocynaceae]]]

Lamiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

38

Plocospermataceae [Oleaceae [Tetrachondraceae [Calceolariaceae

[Gesneriaceae Peltanthera [Pedaliaceae Carlemanniaceae Martyniaceae Stilbaceae

Plantaginaceae Linderniaceae Bignoniaceae Verbenaceae Lamiaceae Paulowniaceae

Schlegeliaceae Thomandersiaceae Phrymaceae Rehmannia Acanthaceae

Scrophulariaceae Orobanchaceae Byblidaceae Lentibulariaceae]]]]]

Solanales

[Montiniaceae [Sphenocleaceae + Hydroleaceae]] [Convolvulaceae + Solanaceae]

CAMPANULIDASTERID II

Aquifoliales

[[Cardiopteridaceae + Stemonuraceae] [Aquifoliaceae [Helwingiaceae +

Phyllonomaceae]]

Asterales

[Rousseaceae + Campanulaceae] Pentaphragmataceae [[Alseuosmiaceae [Phellinaceae

+ Argophyllaceae]] Stylidiaceae [Menyanthaceae [Goodeniaceae [Calyceraceae +

Asteraceae]]]]

Escalloniales

Escalloniaceae

Bruniales

[Bruniaceae + Columelliaceae]

Apiales

Pennantiaceae [Torricelliaceae [Griseliniaceae [Pittosporaceae [Araliaceae

[Myodocarpaceae + Apiaceae]]]]]

Paracryphiales

Paracryphiaceae

Dipsacales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

39

Adoxaceae [Diervillaceae [Caprifoliaceae [Linnaeaceae [Morinaceae [Dipsacaceae +

Valerianaceae]]]]]

LINKS TO ORDERS AND FAMILIES

[Back to Top]

MAIN GROUPINGS

Asterids commelinids core eudicots asterid 1 asterid

2 eudicots gymnosperms Magnoliophyta monocots N-fixing claderosids seed plants

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Abietales Acanthales Acerales Acorales Actinidiales Actinostrobales Adoxales Aescul

ales Agavales Aizoales AkanialesAlismatales Alliales Alseuosmiales Alstroemeriales

Altingiales Amaranthales Amaryllidales Amborellales AmbrosialesAmmiales Amomal

es Ancistrocladales Anisophylleales Annonales Anthobolales Apiales Apocynales Apo

nogetonalesAquifoliales Arales Araliales Aralidiales Araucariales Arecales Aristolochi

ales Athrotaxidales Asarales AsclepiadalesAsparagales Asphodelales Asparagales Ast

eliales Atriplicales Aucubales Austrobaileyales Avenales

Balanitales Balanopales Balanophorales Balsaminales Barbeyales Barclayales Batales

Begoniales BerberidalesBerberidopsidales Betulales Biebersteiniales Bignoniales Bix

ales Boraginales Brassicales Brexiales Bromeliales BrunialesBrunoniales Burmannial

es Burserales Butomales Buxales Byblidales

Cactales Callitrichales Calycanthales Calycerales Campanulales Campynematales Can

ellales Cannales CapparalesCaprifoliales Cardiopteridales Carduales Caricales Carle

manniales Caryophyllales Cassiales Casuarinales CelastralesCentrolepidales Cephalot

ales Cephalotaxales Ceratophyllales Cercidiphyllales Chenopodiales Chironiales Chlor

anthalesChrysobalanales Cinchonales Circaeasterales Cistales Citrales Cocosales Col

chicales Columelliales CombretalesCommelinales Connarales Convolvulales Coriarial

es Cornales Corylales Corynocarpales Crassulales CrossosomatalesCucurbitales Cunn

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

40

inghamiales Cunoniales Cupressales Cycadales Cyclanthales Cymodoceales Cynarales

CynomorialesCyperales Cyrillales Cytinales

Daphnales Daphniphyllales Dasypogonales Datiscales Degeneriales Desfontainiales D

iapensiales Didymelales DillenialesDioales Dioncophyllales Dioscoreales Diospyrales

Dipentodontales Dipsacales Droserales

Ebenales Echiales Elaeagnales Elaeocarpales Elatinales Elodeales Emmotales Empetr

ales Ephedrales Ericales EriocaulalesErythropalaless Escalloniales Eucommiales Euph

orbiales Eupomatiales Eupteleales Euryalaless

Fabales Fagales Falcatifoliales Ficales Flacourtiales Flagellariales Fouquieriales Franc

oales Frangulales

Galiales Garryales Geissolomatales Gentianales Geraniales Gesneriales Ginkgoales G

laucidiales Globulariales GnetalesGoodeniales Greyiales Griseliniales Grossulariales

Grubbiales Gunnerales Gyrocarpales Gyrostemonales

Haemodorales Haloragales Hamamelidales Hanguanales Heisteriales Helleborales He

lwingiales Himantandrales HippuridalesHomaliales Hortensiales Huales Huerteales

Hydatellales Hydnorales Hydrangeales Hydrastidales HydrocharitalesHydropeltidales

Hydrostachyales Hypericales Hypoxidales

Icacinales Illiciales Iridales Irvingales Iteales Ixiales

Jasminales Juglandales Julianiales Juncaginales Juncales

Lacistematales Lactoridales Lamiales Lardizabalales Laurales Lecythidales Ledocarpal

es Leitneriales LentibularialesLigustrales Liliales Limnanthales Linales Liriales Loasal

es Lobeliales Loganiales Lonicerales Loranthales LowialesLythrales

Magnoliales Malpighiales Malvales Marathrales Marcgraviales Mayacales Medusagy

nales Medusandrales MelanthialesMelastomatales Meliales Melianthales Meliosmal

es Menispermales Menyanthales Metteniusales MitrastemonalesMiyoshiales Moni

miales Moringales Musales Myricales Myristicales Myrothamnales Myrsinales Myrta

les

Najadales Nandinales Narcissales Nartheciales Nelumbonales Nepenthales Neuradal

es Nitrariales Nolanales NothofagalesNyctaginales Nymphaeales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

41

Ochnales Oenotherales Olacales Oleales Onagrales Oncothecales Opuntiales Orchid

ales Oxalidales

Saxifragales Pandales Pandanales Papaverales Paracryphiales Parasitaxales Paridales

Parnassiales PassifloralesPenaeales Pennantiales Pentaphragmatales Peridiscales P

etiveriales Petrosaviales Phellinales Philydrales PhyllanthalesPhysenales Phytolaccal

es Picramniales Pinales Pinguiculales Piperales Pittosporales Plantaginales Platanale

sPlumbaginales Poales Podocarpales Podophyllales Podostemales Polemoniales Pol

ygalales Polygonales PontederialesPortulacales Posidoniales Potamogetonales Primu

lales Proteales

Quercales Quillajales Quintiniales

Rafflesiales Ranunculales Rapateales Resedales Restionales Rhabdodendrales Rhamn

ales Rhinanthales RhizophoralesRhodorales Rhoipteleales Roridulales Rosales Rouss

eales Rubiales Ruppiales Rutales

Sabiales Salicales Salvadorales Samolales Samydales Sanguisorbales Santalales Sapin

dales Sapotales SarracenialesSaxegotheales Saxifragales Scheuchzeriales Sciadopitya

les Scleranthales Scrophulariales Scyphostegiales SedalesSilenales Simmondsiales S

milacales Solanales Sphenocleales Sphenostemonales Stangeriales Stellariales Stemo

nalesStilbales Stylidiales Styracales Surianales

Taccales Tamales Tamaricales Taxales Taxodiales Tecophilaeales Terebinthales Tern

stroemiales Theales TheligonalesThymelaeales Tiliales Tofieldiales Torricelliales Tov

ariales Tribelales Trilliales Trimeniales Triuridales TrochodendralesTropaeolales Turn

erales Typhales

Ulmales Urticales

Vacciniales Vahliales Vallisneriales Velloziales Veratrales Verbenales Viburnales Vinc

ales Violales Viscales VitalesVochysiales

Welwitschiales Winterales

Xanthorrhoeales Ximeniales Xyridales

Zamiales Zingiberales Zosterales Zygophyllales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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ALPHABETICAL LISTING OF ALL FAMILY NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | K | L | M | N | O | P | Q | R | S | T | U | V | W | X | Y

|Z

Back to Home

In some cases clicking on a family synonym will send you directly to the subfamily in

which the genus of the basionym is to be found

Abietaceae Abolbodaceae Abrophyllaceae Acalyphaceae Acanthaceae [Acanthochla

mydaceae] Acarnaceaee AceraceaeAchariaceae Achatocarpaceae Achradaceae Achy

ranthaceae Acmopylaceae Acoraceae Acristaceae ActaeaceaeActinidiaceae Actinota

ceae Amentotaxaceae Adenogrammaceae Adoxaceae Aegialitidaceae Aegicerataceae

Aegilopaceae Aeginetiaceaee Aegiphilaceae Aesculaceae Aextoxicaceae Agapanthac

eae Agavaceae AgdestidaceaeAgialidaceae Agrimoniaceae Agrostidaceae Ailanthace

ae Aitoniaceae Aizoaceae Akaniaceae Alangiaceae AlchemillaceaeAldrovandaceae A

lismataceae Alliaceae Allioniaceae Allophylaceae Aloaceae Alopecuraceae Alpiniacea

e AlseuosmiaceaeAlsinaceae Alsodeiaceae Alstroemeriaceae Altingiaceae Alzateacea

e Amaranthaceae Amaryllidaceae Amborellaceae Ambrosiaceae Amentotaxaceae A

mmanniaceae Ammiaceae Amomaceae Ampelidopsaceae Amygdalaceae Amyridacea

eAnacardiaceae Anagallidaceae Anarthriaceae Anchusaceae Ancistrocladaceae Andr

omedaceae AndropogonaceaeAndrostachyaceae Androsynaceae Anemarrhenaceae

Anemonaceae Angelicaceae Anisophylleaceae AnnonaceaeAnomochloaceae Anopter

aceae Anrederaceae Anthemidaceae Anthericaceae Anthobolaceae Antidesmataceae

Antirrhinaceae Antoniaceae Aparinaceae Aphanopetalaceae Aphloiaceae Aphyllanth

aceae Apiaceae ApocynaceaeApodanthaceae Aponogetonaceae Aporusaceae Apose

ridaceae Apostasiaceae Aptandraceae AquifoliaceaeAquilariaceae Aquilegiaceae A

raceae Aragoaceae Araliaceae Aralidiaceaee Araucariaceae Arbutaceae Arceuthidace

aeArceuthobiaceae Arctostaphylaceae Arctotidaceae Ardisiaceae Arecaceae Argoph

yllaceae Arisaraceae AristolochiaceaeAristoteliaceae Arjonaceae Armeriaceae Artem

isiaceae Arthrotaxidaceae Artocarpaceae Arundinaceae ArundinellaceaeAsaraceae A

sclepiadaceae Ascyraceaee Asiraceae Asparagaceae Asperulaceae Asphodelaceae As

pidistraceaeeAsteliaceae Asteraceae Asteranthaceae Asteropeiaceae Astragalaceae

Astrocarpaceae AthanasiaceaeAtherospermataceae Atriplicaceae Atropaceae Aucuba

ceae Aurantiaceae Austrobaileyaceae Austrotaxaceae AvenaceaeAverrhoaceae Avet

raceae Avicenniaceae Azaleaceae Azimaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Balanitaceae Balanopaceae Balanophoraceae Balsameaceae Balsaminaceae Bambusa

ceae Barbaceniaceae BarbeuiaceaeBarbeyaceae Barclayaceae Barringtoniaceae Base

llaceae Bataceae Baueraceae Bauhiniaceae Baxteriaceae BegoniaceaeBehniaceae B

elangeraceae Belloniaceae Bembiciaceae Berberidaceae Berberidopsidaceae Berryac

eaee BersamaceaeBertyaceae Berzeliaceae Besleriaceae Betaceae Betulaceae Biebe

rsteiniaceae Bifariaceae Bignoniaceae BischofiaceaeBixaceae Blakeaceae Blandfordia

ceae Blattiaceae Blepharocaryaceae Blitaceae Blyxaceae Boerlagellaceae Bolivaracea

eBombacaceae Bonnetiaceae Bontiaceae Boopidaceae Boraginaceae Borassaceae B

oroniaceae BoryaceaeBotryodendraceae Bougainvilleaceae Boweniaceae Brassicacea

e Brachycaulaceae Bretschneideraceae BrexiaceaeBromeliaceae Brownlowiaceae Br

unelliaceae Bruniaceae Brunoniaceae Brunsvigiaceae Bryoniaceae BuchneraceaeBuci

daceae Buddlejaceae Buglossaceae Bulbocodiaceae Bumeliaceae Bupleuraceae Burc

hardiaceae BurmanniaceaeBurseraceae Butneriaceae Butomaceae Buxaceae Byblida

ceae Byttneriaceae

Cabombaceae Cacaoaceae Cactaceae Caesalpiniaceae Caladiaceae Calamaceae Calce

olariaceae CalectasiaceaeCalendulaceae Callaceae Callicomaceae Calligonaceae Calli

traceae Callitrichaceae Calochortaceae CalophyllaceaeCalthaceae Calycanthaceae Ca

lyceraceae Cambogiaceae Camelliaceae Campanulaceae CampynemataceaeCanacom

yricaceae Canellaceae Cannabaceae Cannaceae Canopodaceae Canotiaceae Cansjera

ceae CapparaceaeCaprariaceae Caprifoliaceae Cardiopteridaceae Carduaceae Carica

ceae Carissaceae Carlemanniaceae CarpinaceaeCarpodetaceae Cartonemataceae Ca

ryocaraceae Caryophyllaceae Casearieaceae Cassiaceae CassipoureaceaeCassythacea

e Castaneaceae Castelaceae Casuarinaceae Catesbaeaceae Cathedraceae Cecropiace

ae CedraceaeCedrelaceae Celastraceae Celosiaceae Celtidaceae Centaureaceae Cen

trolepidaceae Centroplacaceae CepaceaeCephalanthaceae Cephalotaceae Cephalota

xaceae Cerastiaceae Ceratoniaceae Ceratophyllaceae CerberaceaeCercidiphyllaceae

Cercocarpaceae Cercodiaceae Cereaceae Cerinthaceae Ceroxylaceae Cestraceae Cev

alliaceaeChailletiaceae Chamaedoraceae Chamaemoraceae Chamelauciaceae Chaun

ochitonaceae Chelidoniaceae ChelonaceaeChenopodiaceae Chimonanthaceae Chingi

thamnaceae Chionographidaceae Chiranthodendraceae ChironiaceaeChlaenaceae Ch

loanthaceae Chloranthaceae Chloridaceae Chlorogalaceae Chrysobalanaceae Cicerac

eae CichoriaceaeCimicifugaceae Cinchoniaceae Circaeaceae Circaeasteraceae Cissac

eae Cistaceae Citraceae Clematidaceae CleomaceaeClethraceae Clusiaceae Cneorac

eae Cnestidaceae Cnicaceae Cobaeaceae Cochlospermaceae Cocosaceae Coffeaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Colchicaceae Coleogynaceae Columelliaceae Combretaceae Commelinaceae Comocl

adiaceae Compositae CompsoaceaeConantheraceae Connaraceae Conostylidaceae C

onvallariaceae Convolvulaceae Cordiaceae CoreopsidaceaeCoriandraceae Coriariacea

e Coridaceae Corispermaceae Cornaceae Corokiaceae Coronillaceae Correaceae Cor

rigiolaceaeCorsiaceae Corydalaceae Corylaceae Corynocarpaceae Coryphaceae Cost

aceae Cotyledonaceae CoulaceaeCoutariaceae Coutoubiaceae Crassulaceae Crescen

tiaceae Cressaceae Crinaceae Crocaceae CroomiaceaeCrossosomataceae Crotonacea

e Cruciferae Cryptaceae Crypteroniaceae Cryptocorynaceae CryptomeriaceaeCtenol

ophonaceae Cucurbitaceae Cunninghamiaceae Cunoniaceae Cupressaceae Curcumac

eae Curtisiaceae CuscutaceaeCyananthaceae Cyanastraceae Cynanchaceae Cyanella

ceae Cycadaceae Cyclanthaceae CyclantheraceaeCyclocheilaceae Cydoniaceae Cymo

doceaceae Cynaraceae Cynocrambaceae Cynomoriaceae Cyperaceae CyphiaceaeCyp

hocarpaceae Cypripediaceae Cyrillaceae Cyrtandraceae Cyrtanthaceae Cytinaceae

Dacrycarpaceae Dactylanthaceae Damasoniaceae Daphnaceae Daphniphyllaceae Das

ypogonaceae DatiscaceaeDaturaceae Daucaceae Davidiaceae Davidsoniaceae Decais

neaceae Deeringiaceae Degeneriaceae DelphiniaceaeDendrophthoaceae Columelliac

eae Detariaceae Dialypetalanthaceae Dianellaceae Dianthaceae DiapensiaceaeDicha

petalaceae Dichondraceae Diclidantheraceae Dicrastylidiaceae Dictamnaceae Didiere

aceae DidymelaceaeDidymocarpaceae Diegodendraceae Diervillaceae Digitalidaceae

e Dilatridaceae Dilleniaceae Dioaceae DionaeaceaeDioncophyllaceae Dioscoreaceae

Diosmaceae Diospyraceae Dipentodontaceae Diphylleiaceae Diplolaenaceae Dipsaca

ceaeDipterocarpaceae Dirachmaceae Disanthaceae Diselmaceae Dodonaeaceae Do

mbeyaceae Donatiaceae DorsteniaceaeDortmannaceae Doryanthaceae Drabaceae D

racaenaceae Dracontiaceae Drimyidaceae Droseraceae DrosophyllaceaeDryadaceae

Duabangaceae Duckeodendraceae Dulongiaceae Durantaceae Durionaceae Dysphani

aceae

Ebenaceae Eccremidaceae Ecdeiocoleaceae Echiaceae Echinopaceae Ehretiaceae Ela

eagnaceae ElaeocarpaceaeElegiaceae Elatinaceae Ellisiaceae Ellisiophyllaceae Elodea

ceae Elytranthaceae Embeliaceae Emblingiaceae EmmotaceaeEmpetraceae Encepha

lartaceae Engelhardtiaceae Enhalaceae Epacridaceae Ephedraceae Ephemeranthacea

eEpilobiaceae Epimediaceae Eragrostidaceae Eremolepidaceae Eremosynaceae Erica

ceae Erinaceae EriocaulaceaeEriogonaceae Eriospermaceae Erodiaceae Erycibaceae

Eryngiaceae Erysimaceae Erythroniaceae ErythropalaceaeErythrospermaceae Erythro

xylaceae Escalloniaceae Eschscholziaceae Eucomidaceae Eucommiaceae Eucryphiace

aeEuonymaceae Eupatoriaceae Euphorbiaceae Euphrasiaceae Euphroniaceae Eupo

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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matiaceae Eupteleaceae EuryalaceaeEustrephaceae Euthemidaceae Eutocaceae Exb

ucklandiaceae Exocarpaceae

Fabaceae Fagaceae Falcatifoliaceae Ferulaceae Festucaceae Ficaceae Ficoidaceae Fit

zroyaceae FlacourtiaceaeFlagellariaceae Flindersiaceae Foetidiaceae Forestieraceae

Fothergillaceae Fouquieriaceae Fragariaceae FrancoaceaeFrangulaceae Frankeniacea

e Fraxinaceae Fraxinellaceae Freycinetiaceae Fritillariaceae Fuchsiaceae Fumariacea

eFunkiaceae

Gaiadendraceae Galacaceae Galanthaceae Galaxiaceae Galedupaceae Galeniaceae G

aliaceae Garciniaceae GardeniaceaeGardneriaceae Garryaceae Geissolomataceae Ge

itonoplesiaceae Gelsemiaceae Geniostomaceae GentianaceaeGeonomataceae Geosir

idaceae Geraniaceae Gerrardinaceae Gesneriaceae Gethyllidaceae Gilliesiaceae Gina

lloaceaeGinkgoaceae Gisekiaceae Gladiolaceae Glaucidiaceae Glechomaceae Glinace

ae Globulariaceae Gnaphaliaceae GnetaceaeGoetzeaceae Gomortegaceae Gomphiac

eae Gomphrenaceae Gonystylaceae Goodeniaceae Gordoniaceae GouaniaceaeGoup

iaceae Gramineae Gratiolaceae Grewiaceae Greyiaceae Grielaceae Griseliniaceae Gr

onoviaceae GrossulariaceaeGrubbiaceae Guaiacanaceae Guamatelaceae Guettardac

eae Gunneraceae Gustaviaceae Guttiferae GyrocarpaceaeGyrostemonaceae

Hachetteaceae Haemanthaceae Haemodoraceae Halesiaceae Hallieraceae Halocarpa

ceae Halophilaceae HalophytaceaeHaloragaceae Hamamelidaceae Hanguanaceae H

aptanthaceae Harmandaceae Hebenstretiaceae HectorellaceaeHederaceae Hedyosm

aceae Hedyotidaceae Hedysaraceae Heisteriaceae Heleniaceae Heliamphoraceae H

elianthaceaeHelianthemaceae Helichrysaceae Heliconiaceae Helicteraceae Heliotropi

aceae Helleboraceae Heloniadaceae HelosidaceaeHelwingiaceae Hemerocallidaceae

Hemimeridaceae Hemiphylacaceae Henriqueziaceae Henslowiaceae Hermanniaceae

Hernandiaceae Herniariaceae Herreriaceae Hesperocallidaceae Heterantheraceae He

teropyxidaceae HeterostylaceaeHibbertiaceae Hibiscaceae Hilleriaceae Himantandra

ceae Hippocastanaceae Hippocrateaceae HippomanaceaeHippophaeaceae Hippurida

ceae Hirtellaceae Holacanthaceae Homaliaceae Hopkinsiaceae Hoplestigmataceae H

ordeaceaeHornschuchiaceae Hortensiaceae Hortoniaceae Hostaceae Houstoniaceae

Huaceae Huerteaceae HugoniaceaeHumbertiaceae Humiriaceae Hyacinthaceae Hyd

atellaceae Hydnoraceae Hydrangeaceae Hydrastidaceae HydrillaceaeHydrocharitacea

e Hydrocotylaceae Hydrogetonaceae Hydroleaceae Hydropeltidaceae Hydrophylacac

eae HydrophyllaceaeHydrostachyaceae Hymenocardiaceae Hypecoaceae Hyperanth

eraceae Hypericaceae Hypopityaceae HypoxidaceaeHypseocharitaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta

ceae ImperatoriaceaeInocarpaceae Inulaceae Ionidiaceae Iriarteaceae Iridaceae Irvi

ngiaceae Isnardiaceae Isophysidaceae IteaceaeStrasburgeriaceae Ixiaceae Ixioliriace

ae Ixonanthaceae

Jalapaceae Jamboliferaceae Japonoliriaceae Jasionaceae Jasminiaceae Johnsoniaceae

Joinvilleaceae JuglandaceaeJulianiaceae Juncaceae Juncaginaceae Juniperaceae Jus

siaeaceae Justiciaceae

Kadsuraceae Kaliphoraceae Kalmiaceae Kaniaceae Kiggelariaceae Kingdoniaceae Kin

giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae

Krameriaceae

Labiatae Lacandoniaceae Lachenaliaceae Lacistemataceae Lactoridaceae Lactucaceae

Lagerstroemiaceaee LamiaceaeLampsanaceae Lanariaceae Langsdorffiaceae Lantan

aceae Lapageriaceae Lardizabalaceae Lasiopetalaceae LauraceaeLawsoniaceae Laxm

anniaceae Lecythidaceae Ledaceae Ledocarpaceae Leeaceae Leguminosae Leitneriac

eae LemnaceaeLennoaceae Lentibulariaceae Lentiscaceae Leoniaceae Leonticaceae

Lepidobotryaceae LepidocarpaceaeLepidocaryaceae Lepidocerataceae Leptaulaceae

Leptospermaceae Lepturaceae Lepuropetalaceae LeuctenbergiaceaeLibocedraceae

Licaniaceae Ligustraceae Lilaeaceae Liliaceae Limeaceae Limnanthaceae Limnocharit

aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace

ae Linderniaceae Lindleyaceae Linnaeaceae LippayaceaeLiriaceae Liriodendraceae Li

ssocarpaceae Littorellaceae Loasaceae Lobeliaceaee Loganiaceae LomandraceaeLoni

ceraceae Lopeziaceae Lophiocarpaceae Lophiolaceae Lophiraceae Lophophytaceae L

ophopyxidaceae LoranthaceaeLotaceae Lowiaceae Lupulaceaee Luxemburgiaceae Lu

zuriagaceae Lychnidaceae Lyciaceae LyginiaceaeLygodisodeaceae Lysimachiaceae Ly

thraceae

Macarisiaceae Mackinlayaceae Madiaceae Maesaceae Magnoliaceae Malaceae Male

sherbiaceae MalortieaceaeMalpighiaceae Malvaceae Mangiaceae Manicariaceae Ma

paniaceae Marantaceae Marathraceae MarcgraviaceaeMartyniaceae Mastixiaceae

Matricariaceae Maundiaceae Mayacaceae Medeolaceae Medusagynaceae Peridiscac

eaeMelaleucaceae Melampyraceae Melanophyllaceae Melanthiaceae Melastomatac

eae Meliaceae Melianthaceae MelicaceaeMeliosmaceae Melittidaceae Melochiacea

e Memecylaceae Mendonciaceae Menispermaceae Menthaceae MenyanthaceaeMe

nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

47

Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

Nypaceae Nyssaceae

Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

ceae Papilionaceae Pappophoraceae PapyraceaeParacryphiaceae Parasitaxaceae Par

ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

48

hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

49

cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

50

ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 33: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

33

Arecales

Arecaceae

Poales

[Typhaceae + Bromeliaceae] [Rapateaceae [[Thurniaceae [Juncaceae + Cyperaceae]]

[[Mayacaceae [Eriocaulaceae + Xyridaceae]] [Flagellariaceae [Anarthriaceae

[Centrolepidaceae + Restionaceae]]] [Joinvilleaceae [Ecdeiocoleaceae + Poaceae]]]]]]]

Commelinales

[Commelinaceae + Hanguanaceae] [Philydraceae [Haemodoraceae + Pontederiaceae]]

Zingiberales

Musaceae [Strelitziaceae + Lowiaceae] Heliconiaceae [[Cannaceae + Marantaceae]

[Costaceae + Zingiberaceae]]

Ceratophyllales

Ceratophyllaceae

EUDICOTS

Ranunculales

Eupteleaceae [Papaveraceae [[[Lardizabalaceae + Circaeasteraceae] [Menispermaceae

[Berberidaceae + Ranunculaceae]]]]

Sabiales

Sabiaceae

Proteales

Nelumbonaceae [Platanaceae + Proteaceae]

Trochodendrales

Trochodendraceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

34

Buxales

Haptanthaceae [Buxaceae + Didymelaceae]

CORE EUDICOTS

Gunnerales

Gunneraceae + Myrothamnaceae

Dilleniales

Dilleniaceae

Saxifragales

Peridiscaceae [[Paeoniaceae [Altingiaceae [Hamamelidaceae [Cercidiphyllaceae +

Daphniphyllaceae]]]] [[Crassulaceae [Aphanopetalaceae [Tetracarpaeaceae

[Penthoraceae + Haloragaceae]]]] [Iteaceae [Grossulariaceae + Saxifragaceae]]]]

Cynomoriaceae unplaced

Vitales

Vitaceae

ROSIDS

FABIDROSID I

Zygophyllales

Krameriaceae + Zygophyllaceae

Celastrales

Lepidobotryaceae + Celastraceae

Oxalidales

Huaceae [[Connaraceae + Oxalidaceae] [Cunoniaceae [Elaeocarpaceae [Brunelliaceae +

Cephalotaceae]]]]

Malpighiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

35

[Achariaceae [Goupiaceae [Violaceae + Passifloraceae] [Lacistemataceae Salicaceae]]

[Lophopyxidaceae Putranjivaceae] [Ctenolophonaceae [Erythroxylaceae +

Rhizophoraceae]] Linaceae Ixonanthaceae Humiriaceae Irvingiaceae

Centroplacaceae Pandaceae [[Bonnetiaceae + Clusiaceae] [Calophyllaceae

[Hypericaceae + Podostemaceae]]] [Malpighiaceae + Elatinaceae] Ochnaceae

[Peraceae [Rafflesiaceae + Euphorbiaceae]] [Phyllanthaceae + Picrodendraceae]

[Balanopaceae [[Trigoniaceae + Dichapetalaceae] [Chrysobalanaceae +

Euphroniaceae]]] Caryocaraceae

N-FIXING CLADE

Fabales

Quillajaceae [Fabaceae [Polygalaceae + Surianaceae]]

Rosales

Rosaceae [[Barbeyaceae [Elaeagnaceae Dirachmaceae Rhamnaceae]] [Ulmaceae

[Cannabaceae [Moraceae + Urticaceae]]]]

Cucurbitales

Anisophylleaceae [[Corynocarpaceae + Coriariaceae] [Cucurbitaceae [Tetramelaceae

[Datiscaceae + Begoniaceae]]]] Apodanthaceae

Fagales

Nothofagaceae [Fagaceae [[Myricaceae + Juglandaceae] [Casuarinaceae

[Ticodendraceae + Betulaceae]]]]

MALVIDROSID II

pgtGeraniales

Geraniaceae [[Melianthaceae + Francoaceae] [Vivianaceae + Ledocarpaceae]]

Myrtales

Combretaceae [[Onagraceae + Lythraceae] [[Vochysiaceae + Myrtaceae]

[Melastomataceae [Crypteroniaceae [Alzateaceae + Penaeaceae]]]]]

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

36

Crossosomatales

[Staphyleaceae [Guamatelaceae [Crossosomataceae + Stachyuraceae]]] [Aphloiaceae

[Geissolomataceae + Strasburgeriaceae]]

Picramniales

Picramniaceae

Sapindales

Biebersteiniaceae Nitrariaceae [Kirkiaceae [Anacardiaceae + Burseraceae]]

[Sapindaceae [Simaroubaceae Rutaceae Meliaceae]]

Huerteales

Gerradinaceae [Dipentodontaceae + Tapisciaceae]

Malvales

Neuradaceae [Thymelaeaceae [Sphaerosepalaceae Bixaceae [Cistaceae

[Sarcolaenaceae + Dipterocarpaceae]] [Cytinaceae + Muntingiaceae] Malvaceae]]

Brassicales

[Akaniaceae + Tropaeolaceae] [[Moringaceae + Caricaceae] [Setchellanthaceae

[Limnanthaceae [[Koeberliniaceae [Bataceae + Salvadoraceae]] [Emblingiaceae

[Pentadiplandraceae [Gyrostemonaceae Resedaceae Stixis and relatives] Tovariaceae

[Capparaceae [Cleomaceae + Brassicaceae]]]]]]]]

Berberidopsidales

Aextoxicaceae + Berberidopsidaceae

Santalales

Erythropalaceae Olacaceae [[Loranthaceae [Misodendraceae + Schoepfiaceae]]

[Opiliaceae + Santalaceae]] Balanophoraceae unplaced

Caryophyllales

[[Droseraceae [Nepenthaceae [Drosophyllaceae [Ancistrocladaceae +

Dioncophylleaceae]]]]] [[Frankeniaceae + Tamaricaceae] [Polygonaceae +

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

37

Plumbaginaceae]]] [Rhabdodendraceae [Simmondsiaceae [[Asteropeiaceae +

Physenaceae] [[Caryophyllaceae [Achatocarpaceae + Amaranthaceae]]

[Stegnospermataceae [Limeaceae [[Lophocarpaceae [Barbeuiaceae [Aizoaceae

[Sarcobataceae Phytolaccaceae Nyctaginaceae]]]] [Molluginaceae [Halophytaceae

Montiaceae Basellaceae Didiereaceae [Talinaceae [Portulacaceae [Anacampseros etc +

Cactaceae]]]]]]]]]]]]]

ASTERIDS

Cornales

[Cornaceae + Nyssaceae] [Hydrangeaceae + Loasaceae] Hydrostachyaceae [Curtisiaceae

+ Grubbiaceae]

Ericales

[Balsaminaceae [Marcgraviaceae + Tetrameristaceae]] [[Polemoniaceae +

Fouquieraceae] Lecythidaceae [[Sladeniaceae + Pentaphylacaceae] [Sapotaceae

[Ebenaceae [Maesaceae [Theophrastaceae [Myrsinaceae + Primulaceae]]]]]

[Mitrastemonaceae Theaceae [Symplocaceae [Styracaceae + Diapensiaceae]]

[[Sarraceniaceae [Roridulaceae + Actinidiaceae] [Clethraceae [Cyrillaceae +

Ericaceae]]]]]]

LAMIIDASTERID I

Unplaced

Metteniusaceae Oncothecaceae Icacinaceae and some unplaced ex-Icacinaceae

Garryales

Garryaceae + Eucommiaceae

Unplaced

Boraginaceae Vahliaceae

Gentianales

Rubiaceae [Gentianaceae [Loganiaceae [Gelsemiaceae + Apocynaceae]]]

Lamiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

38

Plocospermataceae [Oleaceae [Tetrachondraceae [Calceolariaceae

[Gesneriaceae Peltanthera [Pedaliaceae Carlemanniaceae Martyniaceae Stilbaceae

Plantaginaceae Linderniaceae Bignoniaceae Verbenaceae Lamiaceae Paulowniaceae

Schlegeliaceae Thomandersiaceae Phrymaceae Rehmannia Acanthaceae

Scrophulariaceae Orobanchaceae Byblidaceae Lentibulariaceae]]]]]

Solanales

[Montiniaceae [Sphenocleaceae + Hydroleaceae]] [Convolvulaceae + Solanaceae]

CAMPANULIDASTERID II

Aquifoliales

[[Cardiopteridaceae + Stemonuraceae] [Aquifoliaceae [Helwingiaceae +

Phyllonomaceae]]

Asterales

[Rousseaceae + Campanulaceae] Pentaphragmataceae [[Alseuosmiaceae [Phellinaceae

+ Argophyllaceae]] Stylidiaceae [Menyanthaceae [Goodeniaceae [Calyceraceae +

Asteraceae]]]]

Escalloniales

Escalloniaceae

Bruniales

[Bruniaceae + Columelliaceae]

Apiales

Pennantiaceae [Torricelliaceae [Griseliniaceae [Pittosporaceae [Araliaceae

[Myodocarpaceae + Apiaceae]]]]]

Paracryphiales

Paracryphiaceae

Dipsacales

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Adoxaceae [Diervillaceae [Caprifoliaceae [Linnaeaceae [Morinaceae [Dipsacaceae +

Valerianaceae]]]]]

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MAIN GROUPINGS

Asterids commelinids core eudicots asterid 1 asterid

2 eudicots gymnosperms Magnoliophyta monocots N-fixing claderosids seed plants

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Abietales Acanthales Acerales Acorales Actinidiales Actinostrobales Adoxales Aescul

ales Agavales Aizoales AkanialesAlismatales Alliales Alseuosmiales Alstroemeriales

Altingiales Amaranthales Amaryllidales Amborellales AmbrosialesAmmiales Amomal

es Ancistrocladales Anisophylleales Annonales Anthobolales Apiales Apocynales Apo

nogetonalesAquifoliales Arales Araliales Aralidiales Araucariales Arecales Aristolochi

ales Athrotaxidales Asarales AsclepiadalesAsparagales Asphodelales Asparagales Ast

eliales Atriplicales Aucubales Austrobaileyales Avenales

Balanitales Balanopales Balanophorales Balsaminales Barbeyales Barclayales Batales

Begoniales BerberidalesBerberidopsidales Betulales Biebersteiniales Bignoniales Bix

ales Boraginales Brassicales Brexiales Bromeliales BrunialesBrunoniales Burmannial

es Burserales Butomales Buxales Byblidales

Cactales Callitrichales Calycanthales Calycerales Campanulales Campynematales Can

ellales Cannales CapparalesCaprifoliales Cardiopteridales Carduales Caricales Carle

manniales Caryophyllales Cassiales Casuarinales CelastralesCentrolepidales Cephalot

ales Cephalotaxales Ceratophyllales Cercidiphyllales Chenopodiales Chironiales Chlor

anthalesChrysobalanales Cinchonales Circaeasterales Cistales Citrales Cocosales Col

chicales Columelliales CombretalesCommelinales Connarales Convolvulales Coriarial

es Cornales Corylales Corynocarpales Crassulales CrossosomatalesCucurbitales Cunn

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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inghamiales Cunoniales Cupressales Cycadales Cyclanthales Cymodoceales Cynarales

CynomorialesCyperales Cyrillales Cytinales

Daphnales Daphniphyllales Dasypogonales Datiscales Degeneriales Desfontainiales D

iapensiales Didymelales DillenialesDioales Dioncophyllales Dioscoreales Diospyrales

Dipentodontales Dipsacales Droserales

Ebenales Echiales Elaeagnales Elaeocarpales Elatinales Elodeales Emmotales Empetr

ales Ephedrales Ericales EriocaulalesErythropalaless Escalloniales Eucommiales Euph

orbiales Eupomatiales Eupteleales Euryalaless

Fabales Fagales Falcatifoliales Ficales Flacourtiales Flagellariales Fouquieriales Franc

oales Frangulales

Galiales Garryales Geissolomatales Gentianales Geraniales Gesneriales Ginkgoales G

laucidiales Globulariales GnetalesGoodeniales Greyiales Griseliniales Grossulariales

Grubbiales Gunnerales Gyrocarpales Gyrostemonales

Haemodorales Haloragales Hamamelidales Hanguanales Heisteriales Helleborales He

lwingiales Himantandrales HippuridalesHomaliales Hortensiales Huales Huerteales

Hydatellales Hydnorales Hydrangeales Hydrastidales HydrocharitalesHydropeltidales

Hydrostachyales Hypericales Hypoxidales

Icacinales Illiciales Iridales Irvingales Iteales Ixiales

Jasminales Juglandales Julianiales Juncaginales Juncales

Lacistematales Lactoridales Lamiales Lardizabalales Laurales Lecythidales Ledocarpal

es Leitneriales LentibularialesLigustrales Liliales Limnanthales Linales Liriales Loasal

es Lobeliales Loganiales Lonicerales Loranthales LowialesLythrales

Magnoliales Malpighiales Malvales Marathrales Marcgraviales Mayacales Medusagy

nales Medusandrales MelanthialesMelastomatales Meliales Melianthales Meliosmal

es Menispermales Menyanthales Metteniusales MitrastemonalesMiyoshiales Moni

miales Moringales Musales Myricales Myristicales Myrothamnales Myrsinales Myrta

les

Najadales Nandinales Narcissales Nartheciales Nelumbonales Nepenthales Neuradal

es Nitrariales Nolanales NothofagalesNyctaginales Nymphaeales

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Ochnales Oenotherales Olacales Oleales Onagrales Oncothecales Opuntiales Orchid

ales Oxalidales

Saxifragales Pandales Pandanales Papaverales Paracryphiales Parasitaxales Paridales

Parnassiales PassifloralesPenaeales Pennantiales Pentaphragmatales Peridiscales P

etiveriales Petrosaviales Phellinales Philydrales PhyllanthalesPhysenales Phytolaccal

es Picramniales Pinales Pinguiculales Piperales Pittosporales Plantaginales Platanale

sPlumbaginales Poales Podocarpales Podophyllales Podostemales Polemoniales Pol

ygalales Polygonales PontederialesPortulacales Posidoniales Potamogetonales Primu

lales Proteales

Quercales Quillajales Quintiniales

Rafflesiales Ranunculales Rapateales Resedales Restionales Rhabdodendrales Rhamn

ales Rhinanthales RhizophoralesRhodorales Rhoipteleales Roridulales Rosales Rouss

eales Rubiales Ruppiales Rutales

Sabiales Salicales Salvadorales Samolales Samydales Sanguisorbales Santalales Sapin

dales Sapotales SarracenialesSaxegotheales Saxifragales Scheuchzeriales Sciadopitya

les Scleranthales Scrophulariales Scyphostegiales SedalesSilenales Simmondsiales S

milacales Solanales Sphenocleales Sphenostemonales Stangeriales Stellariales Stemo

nalesStilbales Stylidiales Styracales Surianales

Taccales Tamales Tamaricales Taxales Taxodiales Tecophilaeales Terebinthales Tern

stroemiales Theales TheligonalesThymelaeales Tiliales Tofieldiales Torricelliales Tov

ariales Tribelales Trilliales Trimeniales Triuridales TrochodendralesTropaeolales Turn

erales Typhales

Ulmales Urticales

Vacciniales Vahliales Vallisneriales Velloziales Veratrales Verbenales Viburnales Vinc

ales Violales Viscales VitalesVochysiales

Welwitschiales Winterales

Xanthorrhoeales Ximeniales Xyridales

Zamiales Zingiberales Zosterales Zygophyllales

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which the genus of the basionym is to be found

Abietaceae Abolbodaceae Abrophyllaceae Acalyphaceae Acanthaceae [Acanthochla

mydaceae] Acarnaceaee AceraceaeAchariaceae Achatocarpaceae Achradaceae Achy

ranthaceae Acmopylaceae Acoraceae Acristaceae ActaeaceaeActinidiaceae Actinota

ceae Amentotaxaceae Adenogrammaceae Adoxaceae Aegialitidaceae Aegicerataceae

Aegilopaceae Aeginetiaceaee Aegiphilaceae Aesculaceae Aextoxicaceae Agapanthac

eae Agavaceae AgdestidaceaeAgialidaceae Agrimoniaceae Agrostidaceae Ailanthace

ae Aitoniaceae Aizoaceae Akaniaceae Alangiaceae AlchemillaceaeAldrovandaceae A

lismataceae Alliaceae Allioniaceae Allophylaceae Aloaceae Alopecuraceae Alpiniacea

e AlseuosmiaceaeAlsinaceae Alsodeiaceae Alstroemeriaceae Altingiaceae Alzateacea

e Amaranthaceae Amaryllidaceae Amborellaceae Ambrosiaceae Amentotaxaceae A

mmanniaceae Ammiaceae Amomaceae Ampelidopsaceae Amygdalaceae Amyridacea

eAnacardiaceae Anagallidaceae Anarthriaceae Anchusaceae Ancistrocladaceae Andr

omedaceae AndropogonaceaeAndrostachyaceae Androsynaceae Anemarrhenaceae

Anemonaceae Angelicaceae Anisophylleaceae AnnonaceaeAnomochloaceae Anopter

aceae Anrederaceae Anthemidaceae Anthericaceae Anthobolaceae Antidesmataceae

Antirrhinaceae Antoniaceae Aparinaceae Aphanopetalaceae Aphloiaceae Aphyllanth

aceae Apiaceae ApocynaceaeApodanthaceae Aponogetonaceae Aporusaceae Apose

ridaceae Apostasiaceae Aptandraceae AquifoliaceaeAquilariaceae Aquilegiaceae A

raceae Aragoaceae Araliaceae Aralidiaceaee Araucariaceae Arbutaceae Arceuthidace

aeArceuthobiaceae Arctostaphylaceae Arctotidaceae Ardisiaceae Arecaceae Argoph

yllaceae Arisaraceae AristolochiaceaeAristoteliaceae Arjonaceae Armeriaceae Artem

isiaceae Arthrotaxidaceae Artocarpaceae Arundinaceae ArundinellaceaeAsaraceae A

sclepiadaceae Ascyraceaee Asiraceae Asparagaceae Asperulaceae Asphodelaceae As

pidistraceaeeAsteliaceae Asteraceae Asteranthaceae Asteropeiaceae Astragalaceae

Astrocarpaceae AthanasiaceaeAtherospermataceae Atriplicaceae Atropaceae Aucuba

ceae Aurantiaceae Austrobaileyaceae Austrotaxaceae AvenaceaeAverrhoaceae Avet

raceae Avicenniaceae Azaleaceae Azimaceae

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Balanitaceae Balanopaceae Balanophoraceae Balsameaceae Balsaminaceae Bambusa

ceae Barbaceniaceae BarbeuiaceaeBarbeyaceae Barclayaceae Barringtoniaceae Base

llaceae Bataceae Baueraceae Bauhiniaceae Baxteriaceae BegoniaceaeBehniaceae B

elangeraceae Belloniaceae Bembiciaceae Berberidaceae Berberidopsidaceae Berryac

eaee BersamaceaeBertyaceae Berzeliaceae Besleriaceae Betaceae Betulaceae Biebe

rsteiniaceae Bifariaceae Bignoniaceae BischofiaceaeBixaceae Blakeaceae Blandfordia

ceae Blattiaceae Blepharocaryaceae Blitaceae Blyxaceae Boerlagellaceae Bolivaracea

eBombacaceae Bonnetiaceae Bontiaceae Boopidaceae Boraginaceae Borassaceae B

oroniaceae BoryaceaeBotryodendraceae Bougainvilleaceae Boweniaceae Brassicacea

e Brachycaulaceae Bretschneideraceae BrexiaceaeBromeliaceae Brownlowiaceae Br

unelliaceae Bruniaceae Brunoniaceae Brunsvigiaceae Bryoniaceae BuchneraceaeBuci

daceae Buddlejaceae Buglossaceae Bulbocodiaceae Bumeliaceae Bupleuraceae Burc

hardiaceae BurmanniaceaeBurseraceae Butneriaceae Butomaceae Buxaceae Byblida

ceae Byttneriaceae

Cabombaceae Cacaoaceae Cactaceae Caesalpiniaceae Caladiaceae Calamaceae Calce

olariaceae CalectasiaceaeCalendulaceae Callaceae Callicomaceae Calligonaceae Calli

traceae Callitrichaceae Calochortaceae CalophyllaceaeCalthaceae Calycanthaceae Ca

lyceraceae Cambogiaceae Camelliaceae Campanulaceae CampynemataceaeCanacom

yricaceae Canellaceae Cannabaceae Cannaceae Canopodaceae Canotiaceae Cansjera

ceae CapparaceaeCaprariaceae Caprifoliaceae Cardiopteridaceae Carduaceae Carica

ceae Carissaceae Carlemanniaceae CarpinaceaeCarpodetaceae Cartonemataceae Ca

ryocaraceae Caryophyllaceae Casearieaceae Cassiaceae CassipoureaceaeCassythacea

e Castaneaceae Castelaceae Casuarinaceae Catesbaeaceae Cathedraceae Cecropiace

ae CedraceaeCedrelaceae Celastraceae Celosiaceae Celtidaceae Centaureaceae Cen

trolepidaceae Centroplacaceae CepaceaeCephalanthaceae Cephalotaceae Cephalota

xaceae Cerastiaceae Ceratoniaceae Ceratophyllaceae CerberaceaeCercidiphyllaceae

Cercocarpaceae Cercodiaceae Cereaceae Cerinthaceae Ceroxylaceae Cestraceae Cev

alliaceaeChailletiaceae Chamaedoraceae Chamaemoraceae Chamelauciaceae Chaun

ochitonaceae Chelidoniaceae ChelonaceaeChenopodiaceae Chimonanthaceae Chingi

thamnaceae Chionographidaceae Chiranthodendraceae ChironiaceaeChlaenaceae Ch

loanthaceae Chloranthaceae Chloridaceae Chlorogalaceae Chrysobalanaceae Cicerac

eae CichoriaceaeCimicifugaceae Cinchoniaceae Circaeaceae Circaeasteraceae Cissac

eae Cistaceae Citraceae Clematidaceae CleomaceaeClethraceae Clusiaceae Cneorac

eae Cnestidaceae Cnicaceae Cobaeaceae Cochlospermaceae Cocosaceae Coffeaceae

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Colchicaceae Coleogynaceae Columelliaceae Combretaceae Commelinaceae Comocl

adiaceae Compositae CompsoaceaeConantheraceae Connaraceae Conostylidaceae C

onvallariaceae Convolvulaceae Cordiaceae CoreopsidaceaeCoriandraceae Coriariacea

e Coridaceae Corispermaceae Cornaceae Corokiaceae Coronillaceae Correaceae Cor

rigiolaceaeCorsiaceae Corydalaceae Corylaceae Corynocarpaceae Coryphaceae Cost

aceae Cotyledonaceae CoulaceaeCoutariaceae Coutoubiaceae Crassulaceae Crescen

tiaceae Cressaceae Crinaceae Crocaceae CroomiaceaeCrossosomataceae Crotonacea

e Cruciferae Cryptaceae Crypteroniaceae Cryptocorynaceae CryptomeriaceaeCtenol

ophonaceae Cucurbitaceae Cunninghamiaceae Cunoniaceae Cupressaceae Curcumac

eae Curtisiaceae CuscutaceaeCyananthaceae Cyanastraceae Cynanchaceae Cyanella

ceae Cycadaceae Cyclanthaceae CyclantheraceaeCyclocheilaceae Cydoniaceae Cymo

doceaceae Cynaraceae Cynocrambaceae Cynomoriaceae Cyperaceae CyphiaceaeCyp

hocarpaceae Cypripediaceae Cyrillaceae Cyrtandraceae Cyrtanthaceae Cytinaceae

Dacrycarpaceae Dactylanthaceae Damasoniaceae Daphnaceae Daphniphyllaceae Das

ypogonaceae DatiscaceaeDaturaceae Daucaceae Davidiaceae Davidsoniaceae Decais

neaceae Deeringiaceae Degeneriaceae DelphiniaceaeDendrophthoaceae Columelliac

eae Detariaceae Dialypetalanthaceae Dianellaceae Dianthaceae DiapensiaceaeDicha

petalaceae Dichondraceae Diclidantheraceae Dicrastylidiaceae Dictamnaceae Didiere

aceae DidymelaceaeDidymocarpaceae Diegodendraceae Diervillaceae Digitalidaceae

e Dilatridaceae Dilleniaceae Dioaceae DionaeaceaeDioncophyllaceae Dioscoreaceae

Diosmaceae Diospyraceae Dipentodontaceae Diphylleiaceae Diplolaenaceae Dipsaca

ceaeDipterocarpaceae Dirachmaceae Disanthaceae Diselmaceae Dodonaeaceae Do

mbeyaceae Donatiaceae DorsteniaceaeDortmannaceae Doryanthaceae Drabaceae D

racaenaceae Dracontiaceae Drimyidaceae Droseraceae DrosophyllaceaeDryadaceae

Duabangaceae Duckeodendraceae Dulongiaceae Durantaceae Durionaceae Dysphani

aceae

Ebenaceae Eccremidaceae Ecdeiocoleaceae Echiaceae Echinopaceae Ehretiaceae Ela

eagnaceae ElaeocarpaceaeElegiaceae Elatinaceae Ellisiaceae Ellisiophyllaceae Elodea

ceae Elytranthaceae Embeliaceae Emblingiaceae EmmotaceaeEmpetraceae Encepha

lartaceae Engelhardtiaceae Enhalaceae Epacridaceae Ephedraceae Ephemeranthacea

eEpilobiaceae Epimediaceae Eragrostidaceae Eremolepidaceae Eremosynaceae Erica

ceae Erinaceae EriocaulaceaeEriogonaceae Eriospermaceae Erodiaceae Erycibaceae

Eryngiaceae Erysimaceae Erythroniaceae ErythropalaceaeErythrospermaceae Erythro

xylaceae Escalloniaceae Eschscholziaceae Eucomidaceae Eucommiaceae Eucryphiace

aeEuonymaceae Eupatoriaceae Euphorbiaceae Euphrasiaceae Euphroniaceae Eupo

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matiaceae Eupteleaceae EuryalaceaeEustrephaceae Euthemidaceae Eutocaceae Exb

ucklandiaceae Exocarpaceae

Fabaceae Fagaceae Falcatifoliaceae Ferulaceae Festucaceae Ficaceae Ficoidaceae Fit

zroyaceae FlacourtiaceaeFlagellariaceae Flindersiaceae Foetidiaceae Forestieraceae

Fothergillaceae Fouquieriaceae Fragariaceae FrancoaceaeFrangulaceae Frankeniacea

e Fraxinaceae Fraxinellaceae Freycinetiaceae Fritillariaceae Fuchsiaceae Fumariacea

eFunkiaceae

Gaiadendraceae Galacaceae Galanthaceae Galaxiaceae Galedupaceae Galeniaceae G

aliaceae Garciniaceae GardeniaceaeGardneriaceae Garryaceae Geissolomataceae Ge

itonoplesiaceae Gelsemiaceae Geniostomaceae GentianaceaeGeonomataceae Geosir

idaceae Geraniaceae Gerrardinaceae Gesneriaceae Gethyllidaceae Gilliesiaceae Gina

lloaceaeGinkgoaceae Gisekiaceae Gladiolaceae Glaucidiaceae Glechomaceae Glinace

ae Globulariaceae Gnaphaliaceae GnetaceaeGoetzeaceae Gomortegaceae Gomphiac

eae Gomphrenaceae Gonystylaceae Goodeniaceae Gordoniaceae GouaniaceaeGoup

iaceae Gramineae Gratiolaceae Grewiaceae Greyiaceae Grielaceae Griseliniaceae Gr

onoviaceae GrossulariaceaeGrubbiaceae Guaiacanaceae Guamatelaceae Guettardac

eae Gunneraceae Gustaviaceae Guttiferae GyrocarpaceaeGyrostemonaceae

Hachetteaceae Haemanthaceae Haemodoraceae Halesiaceae Hallieraceae Halocarpa

ceae Halophilaceae HalophytaceaeHaloragaceae Hamamelidaceae Hanguanaceae H

aptanthaceae Harmandaceae Hebenstretiaceae HectorellaceaeHederaceae Hedyosm

aceae Hedyotidaceae Hedysaraceae Heisteriaceae Heleniaceae Heliamphoraceae H

elianthaceaeHelianthemaceae Helichrysaceae Heliconiaceae Helicteraceae Heliotropi

aceae Helleboraceae Heloniadaceae HelosidaceaeHelwingiaceae Hemerocallidaceae

Hemimeridaceae Hemiphylacaceae Henriqueziaceae Henslowiaceae Hermanniaceae

Hernandiaceae Herniariaceae Herreriaceae Hesperocallidaceae Heterantheraceae He

teropyxidaceae HeterostylaceaeHibbertiaceae Hibiscaceae Hilleriaceae Himantandra

ceae Hippocastanaceae Hippocrateaceae HippomanaceaeHippophaeaceae Hippurida

ceae Hirtellaceae Holacanthaceae Homaliaceae Hopkinsiaceae Hoplestigmataceae H

ordeaceaeHornschuchiaceae Hortensiaceae Hortoniaceae Hostaceae Houstoniaceae

Huaceae Huerteaceae HugoniaceaeHumbertiaceae Humiriaceae Hyacinthaceae Hyd

atellaceae Hydnoraceae Hydrangeaceae Hydrastidaceae HydrillaceaeHydrocharitacea

e Hydrocotylaceae Hydrogetonaceae Hydroleaceae Hydropeltidaceae Hydrophylacac

eae HydrophyllaceaeHydrostachyaceae Hymenocardiaceae Hypecoaceae Hyperanth

eraceae Hypericaceae Hypopityaceae HypoxidaceaeHypseocharitaceae

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Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta

ceae ImperatoriaceaeInocarpaceae Inulaceae Ionidiaceae Iriarteaceae Iridaceae Irvi

ngiaceae Isnardiaceae Isophysidaceae IteaceaeStrasburgeriaceae Ixiaceae Ixioliriace

ae Ixonanthaceae

Jalapaceae Jamboliferaceae Japonoliriaceae Jasionaceae Jasminiaceae Johnsoniaceae

Joinvilleaceae JuglandaceaeJulianiaceae Juncaceae Juncaginaceae Juniperaceae Jus

siaeaceae Justiciaceae

Kadsuraceae Kaliphoraceae Kalmiaceae Kaniaceae Kiggelariaceae Kingdoniaceae Kin

giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae

Krameriaceae

Labiatae Lacandoniaceae Lachenaliaceae Lacistemataceae Lactoridaceae Lactucaceae

Lagerstroemiaceaee LamiaceaeLampsanaceae Lanariaceae Langsdorffiaceae Lantan

aceae Lapageriaceae Lardizabalaceae Lasiopetalaceae LauraceaeLawsoniaceae Laxm

anniaceae Lecythidaceae Ledaceae Ledocarpaceae Leeaceae Leguminosae Leitneriac

eae LemnaceaeLennoaceae Lentibulariaceae Lentiscaceae Leoniaceae Leonticaceae

Lepidobotryaceae LepidocarpaceaeLepidocaryaceae Lepidocerataceae Leptaulaceae

Leptospermaceae Lepturaceae Lepuropetalaceae LeuctenbergiaceaeLibocedraceae

Licaniaceae Ligustraceae Lilaeaceae Liliaceae Limeaceae Limnanthaceae Limnocharit

aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace

ae Linderniaceae Lindleyaceae Linnaeaceae LippayaceaeLiriaceae Liriodendraceae Li

ssocarpaceae Littorellaceae Loasaceae Lobeliaceaee Loganiaceae LomandraceaeLoni

ceraceae Lopeziaceae Lophiocarpaceae Lophiolaceae Lophiraceae Lophophytaceae L

ophopyxidaceae LoranthaceaeLotaceae Lowiaceae Lupulaceaee Luxemburgiaceae Lu

zuriagaceae Lychnidaceae Lyciaceae LyginiaceaeLygodisodeaceae Lysimachiaceae Ly

thraceae

Macarisiaceae Mackinlayaceae Madiaceae Maesaceae Magnoliaceae Malaceae Male

sherbiaceae MalortieaceaeMalpighiaceae Malvaceae Mangiaceae Manicariaceae Ma

paniaceae Marantaceae Marathraceae MarcgraviaceaeMartyniaceae Mastixiaceae

Matricariaceae Maundiaceae Mayacaceae Medeolaceae Medusagynaceae Peridiscac

eaeMelaleucaceae Melampyraceae Melanophyllaceae Melanthiaceae Melastomatac

eae Meliaceae Melianthaceae MelicaceaeMeliosmaceae Melittidaceae Melochiacea

e Memecylaceae Mendonciaceae Menispermaceae Menthaceae MenyanthaceaeMe

nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae

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Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

Nypaceae Nyssaceae

Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

ceae Papilionaceae Pappophoraceae PapyraceaeParacryphiaceae Parasitaxaceae Par

ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

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hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

50

ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 34: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

34

Buxales

Haptanthaceae [Buxaceae + Didymelaceae]

CORE EUDICOTS

Gunnerales

Gunneraceae + Myrothamnaceae

Dilleniales

Dilleniaceae

Saxifragales

Peridiscaceae [[Paeoniaceae [Altingiaceae [Hamamelidaceae [Cercidiphyllaceae +

Daphniphyllaceae]]]] [[Crassulaceae [Aphanopetalaceae [Tetracarpaeaceae

[Penthoraceae + Haloragaceae]]]] [Iteaceae [Grossulariaceae + Saxifragaceae]]]]

Cynomoriaceae unplaced

Vitales

Vitaceae

ROSIDS

FABIDROSID I

Zygophyllales

Krameriaceae + Zygophyllaceae

Celastrales

Lepidobotryaceae + Celastraceae

Oxalidales

Huaceae [[Connaraceae + Oxalidaceae] [Cunoniaceae [Elaeocarpaceae [Brunelliaceae +

Cephalotaceae]]]]

Malpighiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

35

[Achariaceae [Goupiaceae [Violaceae + Passifloraceae] [Lacistemataceae Salicaceae]]

[Lophopyxidaceae Putranjivaceae] [Ctenolophonaceae [Erythroxylaceae +

Rhizophoraceae]] Linaceae Ixonanthaceae Humiriaceae Irvingiaceae

Centroplacaceae Pandaceae [[Bonnetiaceae + Clusiaceae] [Calophyllaceae

[Hypericaceae + Podostemaceae]]] [Malpighiaceae + Elatinaceae] Ochnaceae

[Peraceae [Rafflesiaceae + Euphorbiaceae]] [Phyllanthaceae + Picrodendraceae]

[Balanopaceae [[Trigoniaceae + Dichapetalaceae] [Chrysobalanaceae +

Euphroniaceae]]] Caryocaraceae

N-FIXING CLADE

Fabales

Quillajaceae [Fabaceae [Polygalaceae + Surianaceae]]

Rosales

Rosaceae [[Barbeyaceae [Elaeagnaceae Dirachmaceae Rhamnaceae]] [Ulmaceae

[Cannabaceae [Moraceae + Urticaceae]]]]

Cucurbitales

Anisophylleaceae [[Corynocarpaceae + Coriariaceae] [Cucurbitaceae [Tetramelaceae

[Datiscaceae + Begoniaceae]]]] Apodanthaceae

Fagales

Nothofagaceae [Fagaceae [[Myricaceae + Juglandaceae] [Casuarinaceae

[Ticodendraceae + Betulaceae]]]]

MALVIDROSID II

pgtGeraniales

Geraniaceae [[Melianthaceae + Francoaceae] [Vivianaceae + Ledocarpaceae]]

Myrtales

Combretaceae [[Onagraceae + Lythraceae] [[Vochysiaceae + Myrtaceae]

[Melastomataceae [Crypteroniaceae [Alzateaceae + Penaeaceae]]]]]

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

36

Crossosomatales

[Staphyleaceae [Guamatelaceae [Crossosomataceae + Stachyuraceae]]] [Aphloiaceae

[Geissolomataceae + Strasburgeriaceae]]

Picramniales

Picramniaceae

Sapindales

Biebersteiniaceae Nitrariaceae [Kirkiaceae [Anacardiaceae + Burseraceae]]

[Sapindaceae [Simaroubaceae Rutaceae Meliaceae]]

Huerteales

Gerradinaceae [Dipentodontaceae + Tapisciaceae]

Malvales

Neuradaceae [Thymelaeaceae [Sphaerosepalaceae Bixaceae [Cistaceae

[Sarcolaenaceae + Dipterocarpaceae]] [Cytinaceae + Muntingiaceae] Malvaceae]]

Brassicales

[Akaniaceae + Tropaeolaceae] [[Moringaceae + Caricaceae] [Setchellanthaceae

[Limnanthaceae [[Koeberliniaceae [Bataceae + Salvadoraceae]] [Emblingiaceae

[Pentadiplandraceae [Gyrostemonaceae Resedaceae Stixis and relatives] Tovariaceae

[Capparaceae [Cleomaceae + Brassicaceae]]]]]]]]

Berberidopsidales

Aextoxicaceae + Berberidopsidaceae

Santalales

Erythropalaceae Olacaceae [[Loranthaceae [Misodendraceae + Schoepfiaceae]]

[Opiliaceae + Santalaceae]] Balanophoraceae unplaced

Caryophyllales

[[Droseraceae [Nepenthaceae [Drosophyllaceae [Ancistrocladaceae +

Dioncophylleaceae]]]]] [[Frankeniaceae + Tamaricaceae] [Polygonaceae +

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

37

Plumbaginaceae]]] [Rhabdodendraceae [Simmondsiaceae [[Asteropeiaceae +

Physenaceae] [[Caryophyllaceae [Achatocarpaceae + Amaranthaceae]]

[Stegnospermataceae [Limeaceae [[Lophocarpaceae [Barbeuiaceae [Aizoaceae

[Sarcobataceae Phytolaccaceae Nyctaginaceae]]]] [Molluginaceae [Halophytaceae

Montiaceae Basellaceae Didiereaceae [Talinaceae [Portulacaceae [Anacampseros etc +

Cactaceae]]]]]]]]]]]]]

ASTERIDS

Cornales

[Cornaceae + Nyssaceae] [Hydrangeaceae + Loasaceae] Hydrostachyaceae [Curtisiaceae

+ Grubbiaceae]

Ericales

[Balsaminaceae [Marcgraviaceae + Tetrameristaceae]] [[Polemoniaceae +

Fouquieraceae] Lecythidaceae [[Sladeniaceae + Pentaphylacaceae] [Sapotaceae

[Ebenaceae [Maesaceae [Theophrastaceae [Myrsinaceae + Primulaceae]]]]]

[Mitrastemonaceae Theaceae [Symplocaceae [Styracaceae + Diapensiaceae]]

[[Sarraceniaceae [Roridulaceae + Actinidiaceae] [Clethraceae [Cyrillaceae +

Ericaceae]]]]]]

LAMIIDASTERID I

Unplaced

Metteniusaceae Oncothecaceae Icacinaceae and some unplaced ex-Icacinaceae

Garryales

Garryaceae + Eucommiaceae

Unplaced

Boraginaceae Vahliaceae

Gentianales

Rubiaceae [Gentianaceae [Loganiaceae [Gelsemiaceae + Apocynaceae]]]

Lamiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

38

Plocospermataceae [Oleaceae [Tetrachondraceae [Calceolariaceae

[Gesneriaceae Peltanthera [Pedaliaceae Carlemanniaceae Martyniaceae Stilbaceae

Plantaginaceae Linderniaceae Bignoniaceae Verbenaceae Lamiaceae Paulowniaceae

Schlegeliaceae Thomandersiaceae Phrymaceae Rehmannia Acanthaceae

Scrophulariaceae Orobanchaceae Byblidaceae Lentibulariaceae]]]]]

Solanales

[Montiniaceae [Sphenocleaceae + Hydroleaceae]] [Convolvulaceae + Solanaceae]

CAMPANULIDASTERID II

Aquifoliales

[[Cardiopteridaceae + Stemonuraceae] [Aquifoliaceae [Helwingiaceae +

Phyllonomaceae]]

Asterales

[Rousseaceae + Campanulaceae] Pentaphragmataceae [[Alseuosmiaceae [Phellinaceae

+ Argophyllaceae]] Stylidiaceae [Menyanthaceae [Goodeniaceae [Calyceraceae +

Asteraceae]]]]

Escalloniales

Escalloniaceae

Bruniales

[Bruniaceae + Columelliaceae]

Apiales

Pennantiaceae [Torricelliaceae [Griseliniaceae [Pittosporaceae [Araliaceae

[Myodocarpaceae + Apiaceae]]]]]

Paracryphiales

Paracryphiaceae

Dipsacales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

39

Adoxaceae [Diervillaceae [Caprifoliaceae [Linnaeaceae [Morinaceae [Dipsacaceae +

Valerianaceae]]]]]

LINKS TO ORDERS AND FAMILIES

[Back to Top]

MAIN GROUPINGS

Asterids commelinids core eudicots asterid 1 asterid

2 eudicots gymnosperms Magnoliophyta monocots N-fixing claderosids seed plants

ALPHABETICAL LISTING OF ALL ORDINAL NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | L | M | N | O | P | Q | R | S | T | U | V | W | X | Z

Abietales Acanthales Acerales Acorales Actinidiales Actinostrobales Adoxales Aescul

ales Agavales Aizoales AkanialesAlismatales Alliales Alseuosmiales Alstroemeriales

Altingiales Amaranthales Amaryllidales Amborellales AmbrosialesAmmiales Amomal

es Ancistrocladales Anisophylleales Annonales Anthobolales Apiales Apocynales Apo

nogetonalesAquifoliales Arales Araliales Aralidiales Araucariales Arecales Aristolochi

ales Athrotaxidales Asarales AsclepiadalesAsparagales Asphodelales Asparagales Ast

eliales Atriplicales Aucubales Austrobaileyales Avenales

Balanitales Balanopales Balanophorales Balsaminales Barbeyales Barclayales Batales

Begoniales BerberidalesBerberidopsidales Betulales Biebersteiniales Bignoniales Bix

ales Boraginales Brassicales Brexiales Bromeliales BrunialesBrunoniales Burmannial

es Burserales Butomales Buxales Byblidales

Cactales Callitrichales Calycanthales Calycerales Campanulales Campynematales Can

ellales Cannales CapparalesCaprifoliales Cardiopteridales Carduales Caricales Carle

manniales Caryophyllales Cassiales Casuarinales CelastralesCentrolepidales Cephalot

ales Cephalotaxales Ceratophyllales Cercidiphyllales Chenopodiales Chironiales Chlor

anthalesChrysobalanales Cinchonales Circaeasterales Cistales Citrales Cocosales Col

chicales Columelliales CombretalesCommelinales Connarales Convolvulales Coriarial

es Cornales Corylales Corynocarpales Crassulales CrossosomatalesCucurbitales Cunn

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

40

inghamiales Cunoniales Cupressales Cycadales Cyclanthales Cymodoceales Cynarales

CynomorialesCyperales Cyrillales Cytinales

Daphnales Daphniphyllales Dasypogonales Datiscales Degeneriales Desfontainiales D

iapensiales Didymelales DillenialesDioales Dioncophyllales Dioscoreales Diospyrales

Dipentodontales Dipsacales Droserales

Ebenales Echiales Elaeagnales Elaeocarpales Elatinales Elodeales Emmotales Empetr

ales Ephedrales Ericales EriocaulalesErythropalaless Escalloniales Eucommiales Euph

orbiales Eupomatiales Eupteleales Euryalaless

Fabales Fagales Falcatifoliales Ficales Flacourtiales Flagellariales Fouquieriales Franc

oales Frangulales

Galiales Garryales Geissolomatales Gentianales Geraniales Gesneriales Ginkgoales G

laucidiales Globulariales GnetalesGoodeniales Greyiales Griseliniales Grossulariales

Grubbiales Gunnerales Gyrocarpales Gyrostemonales

Haemodorales Haloragales Hamamelidales Hanguanales Heisteriales Helleborales He

lwingiales Himantandrales HippuridalesHomaliales Hortensiales Huales Huerteales

Hydatellales Hydnorales Hydrangeales Hydrastidales HydrocharitalesHydropeltidales

Hydrostachyales Hypericales Hypoxidales

Icacinales Illiciales Iridales Irvingales Iteales Ixiales

Jasminales Juglandales Julianiales Juncaginales Juncales

Lacistematales Lactoridales Lamiales Lardizabalales Laurales Lecythidales Ledocarpal

es Leitneriales LentibularialesLigustrales Liliales Limnanthales Linales Liriales Loasal

es Lobeliales Loganiales Lonicerales Loranthales LowialesLythrales

Magnoliales Malpighiales Malvales Marathrales Marcgraviales Mayacales Medusagy

nales Medusandrales MelanthialesMelastomatales Meliales Melianthales Meliosmal

es Menispermales Menyanthales Metteniusales MitrastemonalesMiyoshiales Moni

miales Moringales Musales Myricales Myristicales Myrothamnales Myrsinales Myrta

les

Najadales Nandinales Narcissales Nartheciales Nelumbonales Nepenthales Neuradal

es Nitrariales Nolanales NothofagalesNyctaginales Nymphaeales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

41

Ochnales Oenotherales Olacales Oleales Onagrales Oncothecales Opuntiales Orchid

ales Oxalidales

Saxifragales Pandales Pandanales Papaverales Paracryphiales Parasitaxales Paridales

Parnassiales PassifloralesPenaeales Pennantiales Pentaphragmatales Peridiscales P

etiveriales Petrosaviales Phellinales Philydrales PhyllanthalesPhysenales Phytolaccal

es Picramniales Pinales Pinguiculales Piperales Pittosporales Plantaginales Platanale

sPlumbaginales Poales Podocarpales Podophyllales Podostemales Polemoniales Pol

ygalales Polygonales PontederialesPortulacales Posidoniales Potamogetonales Primu

lales Proteales

Quercales Quillajales Quintiniales

Rafflesiales Ranunculales Rapateales Resedales Restionales Rhabdodendrales Rhamn

ales Rhinanthales RhizophoralesRhodorales Rhoipteleales Roridulales Rosales Rouss

eales Rubiales Ruppiales Rutales

Sabiales Salicales Salvadorales Samolales Samydales Sanguisorbales Santalales Sapin

dales Sapotales SarracenialesSaxegotheales Saxifragales Scheuchzeriales Sciadopitya

les Scleranthales Scrophulariales Scyphostegiales SedalesSilenales Simmondsiales S

milacales Solanales Sphenocleales Sphenostemonales Stangeriales Stellariales Stemo

nalesStilbales Stylidiales Styracales Surianales

Taccales Tamales Tamaricales Taxales Taxodiales Tecophilaeales Terebinthales Tern

stroemiales Theales TheligonalesThymelaeales Tiliales Tofieldiales Torricelliales Tov

ariales Tribelales Trilliales Trimeniales Triuridales TrochodendralesTropaeolales Turn

erales Typhales

Ulmales Urticales

Vacciniales Vahliales Vallisneriales Velloziales Veratrales Verbenales Viburnales Vinc

ales Violales Viscales VitalesVochysiales

Welwitschiales Winterales

Xanthorrhoeales Ximeniales Xyridales

Zamiales Zingiberales Zosterales Zygophyllales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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ALPHABETICAL LISTING OF ALL FAMILY NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | K | L | M | N | O | P | Q | R | S | T | U | V | W | X | Y

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In some cases clicking on a family synonym will send you directly to the subfamily in

which the genus of the basionym is to be found

Abietaceae Abolbodaceae Abrophyllaceae Acalyphaceae Acanthaceae [Acanthochla

mydaceae] Acarnaceaee AceraceaeAchariaceae Achatocarpaceae Achradaceae Achy

ranthaceae Acmopylaceae Acoraceae Acristaceae ActaeaceaeActinidiaceae Actinota

ceae Amentotaxaceae Adenogrammaceae Adoxaceae Aegialitidaceae Aegicerataceae

Aegilopaceae Aeginetiaceaee Aegiphilaceae Aesculaceae Aextoxicaceae Agapanthac

eae Agavaceae AgdestidaceaeAgialidaceae Agrimoniaceae Agrostidaceae Ailanthace

ae Aitoniaceae Aizoaceae Akaniaceae Alangiaceae AlchemillaceaeAldrovandaceae A

lismataceae Alliaceae Allioniaceae Allophylaceae Aloaceae Alopecuraceae Alpiniacea

e AlseuosmiaceaeAlsinaceae Alsodeiaceae Alstroemeriaceae Altingiaceae Alzateacea

e Amaranthaceae Amaryllidaceae Amborellaceae Ambrosiaceae Amentotaxaceae A

mmanniaceae Ammiaceae Amomaceae Ampelidopsaceae Amygdalaceae Amyridacea

eAnacardiaceae Anagallidaceae Anarthriaceae Anchusaceae Ancistrocladaceae Andr

omedaceae AndropogonaceaeAndrostachyaceae Androsynaceae Anemarrhenaceae

Anemonaceae Angelicaceae Anisophylleaceae AnnonaceaeAnomochloaceae Anopter

aceae Anrederaceae Anthemidaceae Anthericaceae Anthobolaceae Antidesmataceae

Antirrhinaceae Antoniaceae Aparinaceae Aphanopetalaceae Aphloiaceae Aphyllanth

aceae Apiaceae ApocynaceaeApodanthaceae Aponogetonaceae Aporusaceae Apose

ridaceae Apostasiaceae Aptandraceae AquifoliaceaeAquilariaceae Aquilegiaceae A

raceae Aragoaceae Araliaceae Aralidiaceaee Araucariaceae Arbutaceae Arceuthidace

aeArceuthobiaceae Arctostaphylaceae Arctotidaceae Ardisiaceae Arecaceae Argoph

yllaceae Arisaraceae AristolochiaceaeAristoteliaceae Arjonaceae Armeriaceae Artem

isiaceae Arthrotaxidaceae Artocarpaceae Arundinaceae ArundinellaceaeAsaraceae A

sclepiadaceae Ascyraceaee Asiraceae Asparagaceae Asperulaceae Asphodelaceae As

pidistraceaeeAsteliaceae Asteraceae Asteranthaceae Asteropeiaceae Astragalaceae

Astrocarpaceae AthanasiaceaeAtherospermataceae Atriplicaceae Atropaceae Aucuba

ceae Aurantiaceae Austrobaileyaceae Austrotaxaceae AvenaceaeAverrhoaceae Avet

raceae Avicenniaceae Azaleaceae Azimaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Balanitaceae Balanopaceae Balanophoraceae Balsameaceae Balsaminaceae Bambusa

ceae Barbaceniaceae BarbeuiaceaeBarbeyaceae Barclayaceae Barringtoniaceae Base

llaceae Bataceae Baueraceae Bauhiniaceae Baxteriaceae BegoniaceaeBehniaceae B

elangeraceae Belloniaceae Bembiciaceae Berberidaceae Berberidopsidaceae Berryac

eaee BersamaceaeBertyaceae Berzeliaceae Besleriaceae Betaceae Betulaceae Biebe

rsteiniaceae Bifariaceae Bignoniaceae BischofiaceaeBixaceae Blakeaceae Blandfordia

ceae Blattiaceae Blepharocaryaceae Blitaceae Blyxaceae Boerlagellaceae Bolivaracea

eBombacaceae Bonnetiaceae Bontiaceae Boopidaceae Boraginaceae Borassaceae B

oroniaceae BoryaceaeBotryodendraceae Bougainvilleaceae Boweniaceae Brassicacea

e Brachycaulaceae Bretschneideraceae BrexiaceaeBromeliaceae Brownlowiaceae Br

unelliaceae Bruniaceae Brunoniaceae Brunsvigiaceae Bryoniaceae BuchneraceaeBuci

daceae Buddlejaceae Buglossaceae Bulbocodiaceae Bumeliaceae Bupleuraceae Burc

hardiaceae BurmanniaceaeBurseraceae Butneriaceae Butomaceae Buxaceae Byblida

ceae Byttneriaceae

Cabombaceae Cacaoaceae Cactaceae Caesalpiniaceae Caladiaceae Calamaceae Calce

olariaceae CalectasiaceaeCalendulaceae Callaceae Callicomaceae Calligonaceae Calli

traceae Callitrichaceae Calochortaceae CalophyllaceaeCalthaceae Calycanthaceae Ca

lyceraceae Cambogiaceae Camelliaceae Campanulaceae CampynemataceaeCanacom

yricaceae Canellaceae Cannabaceae Cannaceae Canopodaceae Canotiaceae Cansjera

ceae CapparaceaeCaprariaceae Caprifoliaceae Cardiopteridaceae Carduaceae Carica

ceae Carissaceae Carlemanniaceae CarpinaceaeCarpodetaceae Cartonemataceae Ca

ryocaraceae Caryophyllaceae Casearieaceae Cassiaceae CassipoureaceaeCassythacea

e Castaneaceae Castelaceae Casuarinaceae Catesbaeaceae Cathedraceae Cecropiace

ae CedraceaeCedrelaceae Celastraceae Celosiaceae Celtidaceae Centaureaceae Cen

trolepidaceae Centroplacaceae CepaceaeCephalanthaceae Cephalotaceae Cephalota

xaceae Cerastiaceae Ceratoniaceae Ceratophyllaceae CerberaceaeCercidiphyllaceae

Cercocarpaceae Cercodiaceae Cereaceae Cerinthaceae Ceroxylaceae Cestraceae Cev

alliaceaeChailletiaceae Chamaedoraceae Chamaemoraceae Chamelauciaceae Chaun

ochitonaceae Chelidoniaceae ChelonaceaeChenopodiaceae Chimonanthaceae Chingi

thamnaceae Chionographidaceae Chiranthodendraceae ChironiaceaeChlaenaceae Ch

loanthaceae Chloranthaceae Chloridaceae Chlorogalaceae Chrysobalanaceae Cicerac

eae CichoriaceaeCimicifugaceae Cinchoniaceae Circaeaceae Circaeasteraceae Cissac

eae Cistaceae Citraceae Clematidaceae CleomaceaeClethraceae Clusiaceae Cneorac

eae Cnestidaceae Cnicaceae Cobaeaceae Cochlospermaceae Cocosaceae Coffeaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Colchicaceae Coleogynaceae Columelliaceae Combretaceae Commelinaceae Comocl

adiaceae Compositae CompsoaceaeConantheraceae Connaraceae Conostylidaceae C

onvallariaceae Convolvulaceae Cordiaceae CoreopsidaceaeCoriandraceae Coriariacea

e Coridaceae Corispermaceae Cornaceae Corokiaceae Coronillaceae Correaceae Cor

rigiolaceaeCorsiaceae Corydalaceae Corylaceae Corynocarpaceae Coryphaceae Cost

aceae Cotyledonaceae CoulaceaeCoutariaceae Coutoubiaceae Crassulaceae Crescen

tiaceae Cressaceae Crinaceae Crocaceae CroomiaceaeCrossosomataceae Crotonacea

e Cruciferae Cryptaceae Crypteroniaceae Cryptocorynaceae CryptomeriaceaeCtenol

ophonaceae Cucurbitaceae Cunninghamiaceae Cunoniaceae Cupressaceae Curcumac

eae Curtisiaceae CuscutaceaeCyananthaceae Cyanastraceae Cynanchaceae Cyanella

ceae Cycadaceae Cyclanthaceae CyclantheraceaeCyclocheilaceae Cydoniaceae Cymo

doceaceae Cynaraceae Cynocrambaceae Cynomoriaceae Cyperaceae CyphiaceaeCyp

hocarpaceae Cypripediaceae Cyrillaceae Cyrtandraceae Cyrtanthaceae Cytinaceae

Dacrycarpaceae Dactylanthaceae Damasoniaceae Daphnaceae Daphniphyllaceae Das

ypogonaceae DatiscaceaeDaturaceae Daucaceae Davidiaceae Davidsoniaceae Decais

neaceae Deeringiaceae Degeneriaceae DelphiniaceaeDendrophthoaceae Columelliac

eae Detariaceae Dialypetalanthaceae Dianellaceae Dianthaceae DiapensiaceaeDicha

petalaceae Dichondraceae Diclidantheraceae Dicrastylidiaceae Dictamnaceae Didiere

aceae DidymelaceaeDidymocarpaceae Diegodendraceae Diervillaceae Digitalidaceae

e Dilatridaceae Dilleniaceae Dioaceae DionaeaceaeDioncophyllaceae Dioscoreaceae

Diosmaceae Diospyraceae Dipentodontaceae Diphylleiaceae Diplolaenaceae Dipsaca

ceaeDipterocarpaceae Dirachmaceae Disanthaceae Diselmaceae Dodonaeaceae Do

mbeyaceae Donatiaceae DorsteniaceaeDortmannaceae Doryanthaceae Drabaceae D

racaenaceae Dracontiaceae Drimyidaceae Droseraceae DrosophyllaceaeDryadaceae

Duabangaceae Duckeodendraceae Dulongiaceae Durantaceae Durionaceae Dysphani

aceae

Ebenaceae Eccremidaceae Ecdeiocoleaceae Echiaceae Echinopaceae Ehretiaceae Ela

eagnaceae ElaeocarpaceaeElegiaceae Elatinaceae Ellisiaceae Ellisiophyllaceae Elodea

ceae Elytranthaceae Embeliaceae Emblingiaceae EmmotaceaeEmpetraceae Encepha

lartaceae Engelhardtiaceae Enhalaceae Epacridaceae Ephedraceae Ephemeranthacea

eEpilobiaceae Epimediaceae Eragrostidaceae Eremolepidaceae Eremosynaceae Erica

ceae Erinaceae EriocaulaceaeEriogonaceae Eriospermaceae Erodiaceae Erycibaceae

Eryngiaceae Erysimaceae Erythroniaceae ErythropalaceaeErythrospermaceae Erythro

xylaceae Escalloniaceae Eschscholziaceae Eucomidaceae Eucommiaceae Eucryphiace

aeEuonymaceae Eupatoriaceae Euphorbiaceae Euphrasiaceae Euphroniaceae Eupo

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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matiaceae Eupteleaceae EuryalaceaeEustrephaceae Euthemidaceae Eutocaceae Exb

ucklandiaceae Exocarpaceae

Fabaceae Fagaceae Falcatifoliaceae Ferulaceae Festucaceae Ficaceae Ficoidaceae Fit

zroyaceae FlacourtiaceaeFlagellariaceae Flindersiaceae Foetidiaceae Forestieraceae

Fothergillaceae Fouquieriaceae Fragariaceae FrancoaceaeFrangulaceae Frankeniacea

e Fraxinaceae Fraxinellaceae Freycinetiaceae Fritillariaceae Fuchsiaceae Fumariacea

eFunkiaceae

Gaiadendraceae Galacaceae Galanthaceae Galaxiaceae Galedupaceae Galeniaceae G

aliaceae Garciniaceae GardeniaceaeGardneriaceae Garryaceae Geissolomataceae Ge

itonoplesiaceae Gelsemiaceae Geniostomaceae GentianaceaeGeonomataceae Geosir

idaceae Geraniaceae Gerrardinaceae Gesneriaceae Gethyllidaceae Gilliesiaceae Gina

lloaceaeGinkgoaceae Gisekiaceae Gladiolaceae Glaucidiaceae Glechomaceae Glinace

ae Globulariaceae Gnaphaliaceae GnetaceaeGoetzeaceae Gomortegaceae Gomphiac

eae Gomphrenaceae Gonystylaceae Goodeniaceae Gordoniaceae GouaniaceaeGoup

iaceae Gramineae Gratiolaceae Grewiaceae Greyiaceae Grielaceae Griseliniaceae Gr

onoviaceae GrossulariaceaeGrubbiaceae Guaiacanaceae Guamatelaceae Guettardac

eae Gunneraceae Gustaviaceae Guttiferae GyrocarpaceaeGyrostemonaceae

Hachetteaceae Haemanthaceae Haemodoraceae Halesiaceae Hallieraceae Halocarpa

ceae Halophilaceae HalophytaceaeHaloragaceae Hamamelidaceae Hanguanaceae H

aptanthaceae Harmandaceae Hebenstretiaceae HectorellaceaeHederaceae Hedyosm

aceae Hedyotidaceae Hedysaraceae Heisteriaceae Heleniaceae Heliamphoraceae H

elianthaceaeHelianthemaceae Helichrysaceae Heliconiaceae Helicteraceae Heliotropi

aceae Helleboraceae Heloniadaceae HelosidaceaeHelwingiaceae Hemerocallidaceae

Hemimeridaceae Hemiphylacaceae Henriqueziaceae Henslowiaceae Hermanniaceae

Hernandiaceae Herniariaceae Herreriaceae Hesperocallidaceae Heterantheraceae He

teropyxidaceae HeterostylaceaeHibbertiaceae Hibiscaceae Hilleriaceae Himantandra

ceae Hippocastanaceae Hippocrateaceae HippomanaceaeHippophaeaceae Hippurida

ceae Hirtellaceae Holacanthaceae Homaliaceae Hopkinsiaceae Hoplestigmataceae H

ordeaceaeHornschuchiaceae Hortensiaceae Hortoniaceae Hostaceae Houstoniaceae

Huaceae Huerteaceae HugoniaceaeHumbertiaceae Humiriaceae Hyacinthaceae Hyd

atellaceae Hydnoraceae Hydrangeaceae Hydrastidaceae HydrillaceaeHydrocharitacea

e Hydrocotylaceae Hydrogetonaceae Hydroleaceae Hydropeltidaceae Hydrophylacac

eae HydrophyllaceaeHydrostachyaceae Hymenocardiaceae Hypecoaceae Hyperanth

eraceae Hypericaceae Hypopityaceae HypoxidaceaeHypseocharitaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta

ceae ImperatoriaceaeInocarpaceae Inulaceae Ionidiaceae Iriarteaceae Iridaceae Irvi

ngiaceae Isnardiaceae Isophysidaceae IteaceaeStrasburgeriaceae Ixiaceae Ixioliriace

ae Ixonanthaceae

Jalapaceae Jamboliferaceae Japonoliriaceae Jasionaceae Jasminiaceae Johnsoniaceae

Joinvilleaceae JuglandaceaeJulianiaceae Juncaceae Juncaginaceae Juniperaceae Jus

siaeaceae Justiciaceae

Kadsuraceae Kaliphoraceae Kalmiaceae Kaniaceae Kiggelariaceae Kingdoniaceae Kin

giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae

Krameriaceae

Labiatae Lacandoniaceae Lachenaliaceae Lacistemataceae Lactoridaceae Lactucaceae

Lagerstroemiaceaee LamiaceaeLampsanaceae Lanariaceae Langsdorffiaceae Lantan

aceae Lapageriaceae Lardizabalaceae Lasiopetalaceae LauraceaeLawsoniaceae Laxm

anniaceae Lecythidaceae Ledaceae Ledocarpaceae Leeaceae Leguminosae Leitneriac

eae LemnaceaeLennoaceae Lentibulariaceae Lentiscaceae Leoniaceae Leonticaceae

Lepidobotryaceae LepidocarpaceaeLepidocaryaceae Lepidocerataceae Leptaulaceae

Leptospermaceae Lepturaceae Lepuropetalaceae LeuctenbergiaceaeLibocedraceae

Licaniaceae Ligustraceae Lilaeaceae Liliaceae Limeaceae Limnanthaceae Limnocharit

aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace

ae Linderniaceae Lindleyaceae Linnaeaceae LippayaceaeLiriaceae Liriodendraceae Li

ssocarpaceae Littorellaceae Loasaceae Lobeliaceaee Loganiaceae LomandraceaeLoni

ceraceae Lopeziaceae Lophiocarpaceae Lophiolaceae Lophiraceae Lophophytaceae L

ophopyxidaceae LoranthaceaeLotaceae Lowiaceae Lupulaceaee Luxemburgiaceae Lu

zuriagaceae Lychnidaceae Lyciaceae LyginiaceaeLygodisodeaceae Lysimachiaceae Ly

thraceae

Macarisiaceae Mackinlayaceae Madiaceae Maesaceae Magnoliaceae Malaceae Male

sherbiaceae MalortieaceaeMalpighiaceae Malvaceae Mangiaceae Manicariaceae Ma

paniaceae Marantaceae Marathraceae MarcgraviaceaeMartyniaceae Mastixiaceae

Matricariaceae Maundiaceae Mayacaceae Medeolaceae Medusagynaceae Peridiscac

eaeMelaleucaceae Melampyraceae Melanophyllaceae Melanthiaceae Melastomatac

eae Meliaceae Melianthaceae MelicaceaeMeliosmaceae Melittidaceae Melochiacea

e Memecylaceae Mendonciaceae Menispermaceae Menthaceae MenyanthaceaeMe

nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

Nypaceae Nyssaceae

Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

ceae Papilionaceae Pappophoraceae PapyraceaeParacryphiaceae Parasitaxaceae Par

ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

48

hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

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ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 35: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

35

[Achariaceae [Goupiaceae [Violaceae + Passifloraceae] [Lacistemataceae Salicaceae]]

[Lophopyxidaceae Putranjivaceae] [Ctenolophonaceae [Erythroxylaceae +

Rhizophoraceae]] Linaceae Ixonanthaceae Humiriaceae Irvingiaceae

Centroplacaceae Pandaceae [[Bonnetiaceae + Clusiaceae] [Calophyllaceae

[Hypericaceae + Podostemaceae]]] [Malpighiaceae + Elatinaceae] Ochnaceae

[Peraceae [Rafflesiaceae + Euphorbiaceae]] [Phyllanthaceae + Picrodendraceae]

[Balanopaceae [[Trigoniaceae + Dichapetalaceae] [Chrysobalanaceae +

Euphroniaceae]]] Caryocaraceae

N-FIXING CLADE

Fabales

Quillajaceae [Fabaceae [Polygalaceae + Surianaceae]]

Rosales

Rosaceae [[Barbeyaceae [Elaeagnaceae Dirachmaceae Rhamnaceae]] [Ulmaceae

[Cannabaceae [Moraceae + Urticaceae]]]]

Cucurbitales

Anisophylleaceae [[Corynocarpaceae + Coriariaceae] [Cucurbitaceae [Tetramelaceae

[Datiscaceae + Begoniaceae]]]] Apodanthaceae

Fagales

Nothofagaceae [Fagaceae [[Myricaceae + Juglandaceae] [Casuarinaceae

[Ticodendraceae + Betulaceae]]]]

MALVIDROSID II

pgtGeraniales

Geraniaceae [[Melianthaceae + Francoaceae] [Vivianaceae + Ledocarpaceae]]

Myrtales

Combretaceae [[Onagraceae + Lythraceae] [[Vochysiaceae + Myrtaceae]

[Melastomataceae [Crypteroniaceae [Alzateaceae + Penaeaceae]]]]]

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

36

Crossosomatales

[Staphyleaceae [Guamatelaceae [Crossosomataceae + Stachyuraceae]]] [Aphloiaceae

[Geissolomataceae + Strasburgeriaceae]]

Picramniales

Picramniaceae

Sapindales

Biebersteiniaceae Nitrariaceae [Kirkiaceae [Anacardiaceae + Burseraceae]]

[Sapindaceae [Simaroubaceae Rutaceae Meliaceae]]

Huerteales

Gerradinaceae [Dipentodontaceae + Tapisciaceae]

Malvales

Neuradaceae [Thymelaeaceae [Sphaerosepalaceae Bixaceae [Cistaceae

[Sarcolaenaceae + Dipterocarpaceae]] [Cytinaceae + Muntingiaceae] Malvaceae]]

Brassicales

[Akaniaceae + Tropaeolaceae] [[Moringaceae + Caricaceae] [Setchellanthaceae

[Limnanthaceae [[Koeberliniaceae [Bataceae + Salvadoraceae]] [Emblingiaceae

[Pentadiplandraceae [Gyrostemonaceae Resedaceae Stixis and relatives] Tovariaceae

[Capparaceae [Cleomaceae + Brassicaceae]]]]]]]]

Berberidopsidales

Aextoxicaceae + Berberidopsidaceae

Santalales

Erythropalaceae Olacaceae [[Loranthaceae [Misodendraceae + Schoepfiaceae]]

[Opiliaceae + Santalaceae]] Balanophoraceae unplaced

Caryophyllales

[[Droseraceae [Nepenthaceae [Drosophyllaceae [Ancistrocladaceae +

Dioncophylleaceae]]]]] [[Frankeniaceae + Tamaricaceae] [Polygonaceae +

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

37

Plumbaginaceae]]] [Rhabdodendraceae [Simmondsiaceae [[Asteropeiaceae +

Physenaceae] [[Caryophyllaceae [Achatocarpaceae + Amaranthaceae]]

[Stegnospermataceae [Limeaceae [[Lophocarpaceae [Barbeuiaceae [Aizoaceae

[Sarcobataceae Phytolaccaceae Nyctaginaceae]]]] [Molluginaceae [Halophytaceae

Montiaceae Basellaceae Didiereaceae [Talinaceae [Portulacaceae [Anacampseros etc +

Cactaceae]]]]]]]]]]]]]

ASTERIDS

Cornales

[Cornaceae + Nyssaceae] [Hydrangeaceae + Loasaceae] Hydrostachyaceae [Curtisiaceae

+ Grubbiaceae]

Ericales

[Balsaminaceae [Marcgraviaceae + Tetrameristaceae]] [[Polemoniaceae +

Fouquieraceae] Lecythidaceae [[Sladeniaceae + Pentaphylacaceae] [Sapotaceae

[Ebenaceae [Maesaceae [Theophrastaceae [Myrsinaceae + Primulaceae]]]]]

[Mitrastemonaceae Theaceae [Symplocaceae [Styracaceae + Diapensiaceae]]

[[Sarraceniaceae [Roridulaceae + Actinidiaceae] [Clethraceae [Cyrillaceae +

Ericaceae]]]]]]

LAMIIDASTERID I

Unplaced

Metteniusaceae Oncothecaceae Icacinaceae and some unplaced ex-Icacinaceae

Garryales

Garryaceae + Eucommiaceae

Unplaced

Boraginaceae Vahliaceae

Gentianales

Rubiaceae [Gentianaceae [Loganiaceae [Gelsemiaceae + Apocynaceae]]]

Lamiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

38

Plocospermataceae [Oleaceae [Tetrachondraceae [Calceolariaceae

[Gesneriaceae Peltanthera [Pedaliaceae Carlemanniaceae Martyniaceae Stilbaceae

Plantaginaceae Linderniaceae Bignoniaceae Verbenaceae Lamiaceae Paulowniaceae

Schlegeliaceae Thomandersiaceae Phrymaceae Rehmannia Acanthaceae

Scrophulariaceae Orobanchaceae Byblidaceae Lentibulariaceae]]]]]

Solanales

[Montiniaceae [Sphenocleaceae + Hydroleaceae]] [Convolvulaceae + Solanaceae]

CAMPANULIDASTERID II

Aquifoliales

[[Cardiopteridaceae + Stemonuraceae] [Aquifoliaceae [Helwingiaceae +

Phyllonomaceae]]

Asterales

[Rousseaceae + Campanulaceae] Pentaphragmataceae [[Alseuosmiaceae [Phellinaceae

+ Argophyllaceae]] Stylidiaceae [Menyanthaceae [Goodeniaceae [Calyceraceae +

Asteraceae]]]]

Escalloniales

Escalloniaceae

Bruniales

[Bruniaceae + Columelliaceae]

Apiales

Pennantiaceae [Torricelliaceae [Griseliniaceae [Pittosporaceae [Araliaceae

[Myodocarpaceae + Apiaceae]]]]]

Paracryphiales

Paracryphiaceae

Dipsacales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

39

Adoxaceae [Diervillaceae [Caprifoliaceae [Linnaeaceae [Morinaceae [Dipsacaceae +

Valerianaceae]]]]]

LINKS TO ORDERS AND FAMILIES

[Back to Top]

MAIN GROUPINGS

Asterids commelinids core eudicots asterid 1 asterid

2 eudicots gymnosperms Magnoliophyta monocots N-fixing claderosids seed plants

ALPHABETICAL LISTING OF ALL ORDINAL NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | L | M | N | O | P | Q | R | S | T | U | V | W | X | Z

Abietales Acanthales Acerales Acorales Actinidiales Actinostrobales Adoxales Aescul

ales Agavales Aizoales AkanialesAlismatales Alliales Alseuosmiales Alstroemeriales

Altingiales Amaranthales Amaryllidales Amborellales AmbrosialesAmmiales Amomal

es Ancistrocladales Anisophylleales Annonales Anthobolales Apiales Apocynales Apo

nogetonalesAquifoliales Arales Araliales Aralidiales Araucariales Arecales Aristolochi

ales Athrotaxidales Asarales AsclepiadalesAsparagales Asphodelales Asparagales Ast

eliales Atriplicales Aucubales Austrobaileyales Avenales

Balanitales Balanopales Balanophorales Balsaminales Barbeyales Barclayales Batales

Begoniales BerberidalesBerberidopsidales Betulales Biebersteiniales Bignoniales Bix

ales Boraginales Brassicales Brexiales Bromeliales BrunialesBrunoniales Burmannial

es Burserales Butomales Buxales Byblidales

Cactales Callitrichales Calycanthales Calycerales Campanulales Campynematales Can

ellales Cannales CapparalesCaprifoliales Cardiopteridales Carduales Caricales Carle

manniales Caryophyllales Cassiales Casuarinales CelastralesCentrolepidales Cephalot

ales Cephalotaxales Ceratophyllales Cercidiphyllales Chenopodiales Chironiales Chlor

anthalesChrysobalanales Cinchonales Circaeasterales Cistales Citrales Cocosales Col

chicales Columelliales CombretalesCommelinales Connarales Convolvulales Coriarial

es Cornales Corylales Corynocarpales Crassulales CrossosomatalesCucurbitales Cunn

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

40

inghamiales Cunoniales Cupressales Cycadales Cyclanthales Cymodoceales Cynarales

CynomorialesCyperales Cyrillales Cytinales

Daphnales Daphniphyllales Dasypogonales Datiscales Degeneriales Desfontainiales D

iapensiales Didymelales DillenialesDioales Dioncophyllales Dioscoreales Diospyrales

Dipentodontales Dipsacales Droserales

Ebenales Echiales Elaeagnales Elaeocarpales Elatinales Elodeales Emmotales Empetr

ales Ephedrales Ericales EriocaulalesErythropalaless Escalloniales Eucommiales Euph

orbiales Eupomatiales Eupteleales Euryalaless

Fabales Fagales Falcatifoliales Ficales Flacourtiales Flagellariales Fouquieriales Franc

oales Frangulales

Galiales Garryales Geissolomatales Gentianales Geraniales Gesneriales Ginkgoales G

laucidiales Globulariales GnetalesGoodeniales Greyiales Griseliniales Grossulariales

Grubbiales Gunnerales Gyrocarpales Gyrostemonales

Haemodorales Haloragales Hamamelidales Hanguanales Heisteriales Helleborales He

lwingiales Himantandrales HippuridalesHomaliales Hortensiales Huales Huerteales

Hydatellales Hydnorales Hydrangeales Hydrastidales HydrocharitalesHydropeltidales

Hydrostachyales Hypericales Hypoxidales

Icacinales Illiciales Iridales Irvingales Iteales Ixiales

Jasminales Juglandales Julianiales Juncaginales Juncales

Lacistematales Lactoridales Lamiales Lardizabalales Laurales Lecythidales Ledocarpal

es Leitneriales LentibularialesLigustrales Liliales Limnanthales Linales Liriales Loasal

es Lobeliales Loganiales Lonicerales Loranthales LowialesLythrales

Magnoliales Malpighiales Malvales Marathrales Marcgraviales Mayacales Medusagy

nales Medusandrales MelanthialesMelastomatales Meliales Melianthales Meliosmal

es Menispermales Menyanthales Metteniusales MitrastemonalesMiyoshiales Moni

miales Moringales Musales Myricales Myristicales Myrothamnales Myrsinales Myrta

les

Najadales Nandinales Narcissales Nartheciales Nelumbonales Nepenthales Neuradal

es Nitrariales Nolanales NothofagalesNyctaginales Nymphaeales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

41

Ochnales Oenotherales Olacales Oleales Onagrales Oncothecales Opuntiales Orchid

ales Oxalidales

Saxifragales Pandales Pandanales Papaverales Paracryphiales Parasitaxales Paridales

Parnassiales PassifloralesPenaeales Pennantiales Pentaphragmatales Peridiscales P

etiveriales Petrosaviales Phellinales Philydrales PhyllanthalesPhysenales Phytolaccal

es Picramniales Pinales Pinguiculales Piperales Pittosporales Plantaginales Platanale

sPlumbaginales Poales Podocarpales Podophyllales Podostemales Polemoniales Pol

ygalales Polygonales PontederialesPortulacales Posidoniales Potamogetonales Primu

lales Proteales

Quercales Quillajales Quintiniales

Rafflesiales Ranunculales Rapateales Resedales Restionales Rhabdodendrales Rhamn

ales Rhinanthales RhizophoralesRhodorales Rhoipteleales Roridulales Rosales Rouss

eales Rubiales Ruppiales Rutales

Sabiales Salicales Salvadorales Samolales Samydales Sanguisorbales Santalales Sapin

dales Sapotales SarracenialesSaxegotheales Saxifragales Scheuchzeriales Sciadopitya

les Scleranthales Scrophulariales Scyphostegiales SedalesSilenales Simmondsiales S

milacales Solanales Sphenocleales Sphenostemonales Stangeriales Stellariales Stemo

nalesStilbales Stylidiales Styracales Surianales

Taccales Tamales Tamaricales Taxales Taxodiales Tecophilaeales Terebinthales Tern

stroemiales Theales TheligonalesThymelaeales Tiliales Tofieldiales Torricelliales Tov

ariales Tribelales Trilliales Trimeniales Triuridales TrochodendralesTropaeolales Turn

erales Typhales

Ulmales Urticales

Vacciniales Vahliales Vallisneriales Velloziales Veratrales Verbenales Viburnales Vinc

ales Violales Viscales VitalesVochysiales

Welwitschiales Winterales

Xanthorrhoeales Ximeniales Xyridales

Zamiales Zingiberales Zosterales Zygophyllales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

42

ALPHABETICAL LISTING OF ALL FAMILY NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | K | L | M | N | O | P | Q | R | S | T | U | V | W | X | Y

|Z

Back to Home

In some cases clicking on a family synonym will send you directly to the subfamily in

which the genus of the basionym is to be found

Abietaceae Abolbodaceae Abrophyllaceae Acalyphaceae Acanthaceae [Acanthochla

mydaceae] Acarnaceaee AceraceaeAchariaceae Achatocarpaceae Achradaceae Achy

ranthaceae Acmopylaceae Acoraceae Acristaceae ActaeaceaeActinidiaceae Actinota

ceae Amentotaxaceae Adenogrammaceae Adoxaceae Aegialitidaceae Aegicerataceae

Aegilopaceae Aeginetiaceaee Aegiphilaceae Aesculaceae Aextoxicaceae Agapanthac

eae Agavaceae AgdestidaceaeAgialidaceae Agrimoniaceae Agrostidaceae Ailanthace

ae Aitoniaceae Aizoaceae Akaniaceae Alangiaceae AlchemillaceaeAldrovandaceae A

lismataceae Alliaceae Allioniaceae Allophylaceae Aloaceae Alopecuraceae Alpiniacea

e AlseuosmiaceaeAlsinaceae Alsodeiaceae Alstroemeriaceae Altingiaceae Alzateacea

e Amaranthaceae Amaryllidaceae Amborellaceae Ambrosiaceae Amentotaxaceae A

mmanniaceae Ammiaceae Amomaceae Ampelidopsaceae Amygdalaceae Amyridacea

eAnacardiaceae Anagallidaceae Anarthriaceae Anchusaceae Ancistrocladaceae Andr

omedaceae AndropogonaceaeAndrostachyaceae Androsynaceae Anemarrhenaceae

Anemonaceae Angelicaceae Anisophylleaceae AnnonaceaeAnomochloaceae Anopter

aceae Anrederaceae Anthemidaceae Anthericaceae Anthobolaceae Antidesmataceae

Antirrhinaceae Antoniaceae Aparinaceae Aphanopetalaceae Aphloiaceae Aphyllanth

aceae Apiaceae ApocynaceaeApodanthaceae Aponogetonaceae Aporusaceae Apose

ridaceae Apostasiaceae Aptandraceae AquifoliaceaeAquilariaceae Aquilegiaceae A

raceae Aragoaceae Araliaceae Aralidiaceaee Araucariaceae Arbutaceae Arceuthidace

aeArceuthobiaceae Arctostaphylaceae Arctotidaceae Ardisiaceae Arecaceae Argoph

yllaceae Arisaraceae AristolochiaceaeAristoteliaceae Arjonaceae Armeriaceae Artem

isiaceae Arthrotaxidaceae Artocarpaceae Arundinaceae ArundinellaceaeAsaraceae A

sclepiadaceae Ascyraceaee Asiraceae Asparagaceae Asperulaceae Asphodelaceae As

pidistraceaeeAsteliaceae Asteraceae Asteranthaceae Asteropeiaceae Astragalaceae

Astrocarpaceae AthanasiaceaeAtherospermataceae Atriplicaceae Atropaceae Aucuba

ceae Aurantiaceae Austrobaileyaceae Austrotaxaceae AvenaceaeAverrhoaceae Avet

raceae Avicenniaceae Azaleaceae Azimaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

43

Balanitaceae Balanopaceae Balanophoraceae Balsameaceae Balsaminaceae Bambusa

ceae Barbaceniaceae BarbeuiaceaeBarbeyaceae Barclayaceae Barringtoniaceae Base

llaceae Bataceae Baueraceae Bauhiniaceae Baxteriaceae BegoniaceaeBehniaceae B

elangeraceae Belloniaceae Bembiciaceae Berberidaceae Berberidopsidaceae Berryac

eaee BersamaceaeBertyaceae Berzeliaceae Besleriaceae Betaceae Betulaceae Biebe

rsteiniaceae Bifariaceae Bignoniaceae BischofiaceaeBixaceae Blakeaceae Blandfordia

ceae Blattiaceae Blepharocaryaceae Blitaceae Blyxaceae Boerlagellaceae Bolivaracea

eBombacaceae Bonnetiaceae Bontiaceae Boopidaceae Boraginaceae Borassaceae B

oroniaceae BoryaceaeBotryodendraceae Bougainvilleaceae Boweniaceae Brassicacea

e Brachycaulaceae Bretschneideraceae BrexiaceaeBromeliaceae Brownlowiaceae Br

unelliaceae Bruniaceae Brunoniaceae Brunsvigiaceae Bryoniaceae BuchneraceaeBuci

daceae Buddlejaceae Buglossaceae Bulbocodiaceae Bumeliaceae Bupleuraceae Burc

hardiaceae BurmanniaceaeBurseraceae Butneriaceae Butomaceae Buxaceae Byblida

ceae Byttneriaceae

Cabombaceae Cacaoaceae Cactaceae Caesalpiniaceae Caladiaceae Calamaceae Calce

olariaceae CalectasiaceaeCalendulaceae Callaceae Callicomaceae Calligonaceae Calli

traceae Callitrichaceae Calochortaceae CalophyllaceaeCalthaceae Calycanthaceae Ca

lyceraceae Cambogiaceae Camelliaceae Campanulaceae CampynemataceaeCanacom

yricaceae Canellaceae Cannabaceae Cannaceae Canopodaceae Canotiaceae Cansjera

ceae CapparaceaeCaprariaceae Caprifoliaceae Cardiopteridaceae Carduaceae Carica

ceae Carissaceae Carlemanniaceae CarpinaceaeCarpodetaceae Cartonemataceae Ca

ryocaraceae Caryophyllaceae Casearieaceae Cassiaceae CassipoureaceaeCassythacea

e Castaneaceae Castelaceae Casuarinaceae Catesbaeaceae Cathedraceae Cecropiace

ae CedraceaeCedrelaceae Celastraceae Celosiaceae Celtidaceae Centaureaceae Cen

trolepidaceae Centroplacaceae CepaceaeCephalanthaceae Cephalotaceae Cephalota

xaceae Cerastiaceae Ceratoniaceae Ceratophyllaceae CerberaceaeCercidiphyllaceae

Cercocarpaceae Cercodiaceae Cereaceae Cerinthaceae Ceroxylaceae Cestraceae Cev

alliaceaeChailletiaceae Chamaedoraceae Chamaemoraceae Chamelauciaceae Chaun

ochitonaceae Chelidoniaceae ChelonaceaeChenopodiaceae Chimonanthaceae Chingi

thamnaceae Chionographidaceae Chiranthodendraceae ChironiaceaeChlaenaceae Ch

loanthaceae Chloranthaceae Chloridaceae Chlorogalaceae Chrysobalanaceae Cicerac

eae CichoriaceaeCimicifugaceae Cinchoniaceae Circaeaceae Circaeasteraceae Cissac

eae Cistaceae Citraceae Clematidaceae CleomaceaeClethraceae Clusiaceae Cneorac

eae Cnestidaceae Cnicaceae Cobaeaceae Cochlospermaceae Cocosaceae Coffeaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

44

Colchicaceae Coleogynaceae Columelliaceae Combretaceae Commelinaceae Comocl

adiaceae Compositae CompsoaceaeConantheraceae Connaraceae Conostylidaceae C

onvallariaceae Convolvulaceae Cordiaceae CoreopsidaceaeCoriandraceae Coriariacea

e Coridaceae Corispermaceae Cornaceae Corokiaceae Coronillaceae Correaceae Cor

rigiolaceaeCorsiaceae Corydalaceae Corylaceae Corynocarpaceae Coryphaceae Cost

aceae Cotyledonaceae CoulaceaeCoutariaceae Coutoubiaceae Crassulaceae Crescen

tiaceae Cressaceae Crinaceae Crocaceae CroomiaceaeCrossosomataceae Crotonacea

e Cruciferae Cryptaceae Crypteroniaceae Cryptocorynaceae CryptomeriaceaeCtenol

ophonaceae Cucurbitaceae Cunninghamiaceae Cunoniaceae Cupressaceae Curcumac

eae Curtisiaceae CuscutaceaeCyananthaceae Cyanastraceae Cynanchaceae Cyanella

ceae Cycadaceae Cyclanthaceae CyclantheraceaeCyclocheilaceae Cydoniaceae Cymo

doceaceae Cynaraceae Cynocrambaceae Cynomoriaceae Cyperaceae CyphiaceaeCyp

hocarpaceae Cypripediaceae Cyrillaceae Cyrtandraceae Cyrtanthaceae Cytinaceae

Dacrycarpaceae Dactylanthaceae Damasoniaceae Daphnaceae Daphniphyllaceae Das

ypogonaceae DatiscaceaeDaturaceae Daucaceae Davidiaceae Davidsoniaceae Decais

neaceae Deeringiaceae Degeneriaceae DelphiniaceaeDendrophthoaceae Columelliac

eae Detariaceae Dialypetalanthaceae Dianellaceae Dianthaceae DiapensiaceaeDicha

petalaceae Dichondraceae Diclidantheraceae Dicrastylidiaceae Dictamnaceae Didiere

aceae DidymelaceaeDidymocarpaceae Diegodendraceae Diervillaceae Digitalidaceae

e Dilatridaceae Dilleniaceae Dioaceae DionaeaceaeDioncophyllaceae Dioscoreaceae

Diosmaceae Diospyraceae Dipentodontaceae Diphylleiaceae Diplolaenaceae Dipsaca

ceaeDipterocarpaceae Dirachmaceae Disanthaceae Diselmaceae Dodonaeaceae Do

mbeyaceae Donatiaceae DorsteniaceaeDortmannaceae Doryanthaceae Drabaceae D

racaenaceae Dracontiaceae Drimyidaceae Droseraceae DrosophyllaceaeDryadaceae

Duabangaceae Duckeodendraceae Dulongiaceae Durantaceae Durionaceae Dysphani

aceae

Ebenaceae Eccremidaceae Ecdeiocoleaceae Echiaceae Echinopaceae Ehretiaceae Ela

eagnaceae ElaeocarpaceaeElegiaceae Elatinaceae Ellisiaceae Ellisiophyllaceae Elodea

ceae Elytranthaceae Embeliaceae Emblingiaceae EmmotaceaeEmpetraceae Encepha

lartaceae Engelhardtiaceae Enhalaceae Epacridaceae Ephedraceae Ephemeranthacea

eEpilobiaceae Epimediaceae Eragrostidaceae Eremolepidaceae Eremosynaceae Erica

ceae Erinaceae EriocaulaceaeEriogonaceae Eriospermaceae Erodiaceae Erycibaceae

Eryngiaceae Erysimaceae Erythroniaceae ErythropalaceaeErythrospermaceae Erythro

xylaceae Escalloniaceae Eschscholziaceae Eucomidaceae Eucommiaceae Eucryphiace

aeEuonymaceae Eupatoriaceae Euphorbiaceae Euphrasiaceae Euphroniaceae Eupo

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

45

matiaceae Eupteleaceae EuryalaceaeEustrephaceae Euthemidaceae Eutocaceae Exb

ucklandiaceae Exocarpaceae

Fabaceae Fagaceae Falcatifoliaceae Ferulaceae Festucaceae Ficaceae Ficoidaceae Fit

zroyaceae FlacourtiaceaeFlagellariaceae Flindersiaceae Foetidiaceae Forestieraceae

Fothergillaceae Fouquieriaceae Fragariaceae FrancoaceaeFrangulaceae Frankeniacea

e Fraxinaceae Fraxinellaceae Freycinetiaceae Fritillariaceae Fuchsiaceae Fumariacea

eFunkiaceae

Gaiadendraceae Galacaceae Galanthaceae Galaxiaceae Galedupaceae Galeniaceae G

aliaceae Garciniaceae GardeniaceaeGardneriaceae Garryaceae Geissolomataceae Ge

itonoplesiaceae Gelsemiaceae Geniostomaceae GentianaceaeGeonomataceae Geosir

idaceae Geraniaceae Gerrardinaceae Gesneriaceae Gethyllidaceae Gilliesiaceae Gina

lloaceaeGinkgoaceae Gisekiaceae Gladiolaceae Glaucidiaceae Glechomaceae Glinace

ae Globulariaceae Gnaphaliaceae GnetaceaeGoetzeaceae Gomortegaceae Gomphiac

eae Gomphrenaceae Gonystylaceae Goodeniaceae Gordoniaceae GouaniaceaeGoup

iaceae Gramineae Gratiolaceae Grewiaceae Greyiaceae Grielaceae Griseliniaceae Gr

onoviaceae GrossulariaceaeGrubbiaceae Guaiacanaceae Guamatelaceae Guettardac

eae Gunneraceae Gustaviaceae Guttiferae GyrocarpaceaeGyrostemonaceae

Hachetteaceae Haemanthaceae Haemodoraceae Halesiaceae Hallieraceae Halocarpa

ceae Halophilaceae HalophytaceaeHaloragaceae Hamamelidaceae Hanguanaceae H

aptanthaceae Harmandaceae Hebenstretiaceae HectorellaceaeHederaceae Hedyosm

aceae Hedyotidaceae Hedysaraceae Heisteriaceae Heleniaceae Heliamphoraceae H

elianthaceaeHelianthemaceae Helichrysaceae Heliconiaceae Helicteraceae Heliotropi

aceae Helleboraceae Heloniadaceae HelosidaceaeHelwingiaceae Hemerocallidaceae

Hemimeridaceae Hemiphylacaceae Henriqueziaceae Henslowiaceae Hermanniaceae

Hernandiaceae Herniariaceae Herreriaceae Hesperocallidaceae Heterantheraceae He

teropyxidaceae HeterostylaceaeHibbertiaceae Hibiscaceae Hilleriaceae Himantandra

ceae Hippocastanaceae Hippocrateaceae HippomanaceaeHippophaeaceae Hippurida

ceae Hirtellaceae Holacanthaceae Homaliaceae Hopkinsiaceae Hoplestigmataceae H

ordeaceaeHornschuchiaceae Hortensiaceae Hortoniaceae Hostaceae Houstoniaceae

Huaceae Huerteaceae HugoniaceaeHumbertiaceae Humiriaceae Hyacinthaceae Hyd

atellaceae Hydnoraceae Hydrangeaceae Hydrastidaceae HydrillaceaeHydrocharitacea

e Hydrocotylaceae Hydrogetonaceae Hydroleaceae Hydropeltidaceae Hydrophylacac

eae HydrophyllaceaeHydrostachyaceae Hymenocardiaceae Hypecoaceae Hyperanth

eraceae Hypericaceae Hypopityaceae HypoxidaceaeHypseocharitaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta

ceae ImperatoriaceaeInocarpaceae Inulaceae Ionidiaceae Iriarteaceae Iridaceae Irvi

ngiaceae Isnardiaceae Isophysidaceae IteaceaeStrasburgeriaceae Ixiaceae Ixioliriace

ae Ixonanthaceae

Jalapaceae Jamboliferaceae Japonoliriaceae Jasionaceae Jasminiaceae Johnsoniaceae

Joinvilleaceae JuglandaceaeJulianiaceae Juncaceae Juncaginaceae Juniperaceae Jus

siaeaceae Justiciaceae

Kadsuraceae Kaliphoraceae Kalmiaceae Kaniaceae Kiggelariaceae Kingdoniaceae Kin

giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae

Krameriaceae

Labiatae Lacandoniaceae Lachenaliaceae Lacistemataceae Lactoridaceae Lactucaceae

Lagerstroemiaceaee LamiaceaeLampsanaceae Lanariaceae Langsdorffiaceae Lantan

aceae Lapageriaceae Lardizabalaceae Lasiopetalaceae LauraceaeLawsoniaceae Laxm

anniaceae Lecythidaceae Ledaceae Ledocarpaceae Leeaceae Leguminosae Leitneriac

eae LemnaceaeLennoaceae Lentibulariaceae Lentiscaceae Leoniaceae Leonticaceae

Lepidobotryaceae LepidocarpaceaeLepidocaryaceae Lepidocerataceae Leptaulaceae

Leptospermaceae Lepturaceae Lepuropetalaceae LeuctenbergiaceaeLibocedraceae

Licaniaceae Ligustraceae Lilaeaceae Liliaceae Limeaceae Limnanthaceae Limnocharit

aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace

ae Linderniaceae Lindleyaceae Linnaeaceae LippayaceaeLiriaceae Liriodendraceae Li

ssocarpaceae Littorellaceae Loasaceae Lobeliaceaee Loganiaceae LomandraceaeLoni

ceraceae Lopeziaceae Lophiocarpaceae Lophiolaceae Lophiraceae Lophophytaceae L

ophopyxidaceae LoranthaceaeLotaceae Lowiaceae Lupulaceaee Luxemburgiaceae Lu

zuriagaceae Lychnidaceae Lyciaceae LyginiaceaeLygodisodeaceae Lysimachiaceae Ly

thraceae

Macarisiaceae Mackinlayaceae Madiaceae Maesaceae Magnoliaceae Malaceae Male

sherbiaceae MalortieaceaeMalpighiaceae Malvaceae Mangiaceae Manicariaceae Ma

paniaceae Marantaceae Marathraceae MarcgraviaceaeMartyniaceae Mastixiaceae

Matricariaceae Maundiaceae Mayacaceae Medeolaceae Medusagynaceae Peridiscac

eaeMelaleucaceae Melampyraceae Melanophyllaceae Melanthiaceae Melastomatac

eae Meliaceae Melianthaceae MelicaceaeMeliosmaceae Melittidaceae Melochiacea

e Memecylaceae Mendonciaceae Menispermaceae Menthaceae MenyanthaceaeMe

nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

47

Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

Nypaceae Nyssaceae

Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

ceae Papilionaceae Pappophoraceae PapyraceaeParacryphiaceae Parasitaxaceae Par

ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

48

hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

49

cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

50

ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 36: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

36

Crossosomatales

[Staphyleaceae [Guamatelaceae [Crossosomataceae + Stachyuraceae]]] [Aphloiaceae

[Geissolomataceae + Strasburgeriaceae]]

Picramniales

Picramniaceae

Sapindales

Biebersteiniaceae Nitrariaceae [Kirkiaceae [Anacardiaceae + Burseraceae]]

[Sapindaceae [Simaroubaceae Rutaceae Meliaceae]]

Huerteales

Gerradinaceae [Dipentodontaceae + Tapisciaceae]

Malvales

Neuradaceae [Thymelaeaceae [Sphaerosepalaceae Bixaceae [Cistaceae

[Sarcolaenaceae + Dipterocarpaceae]] [Cytinaceae + Muntingiaceae] Malvaceae]]

Brassicales

[Akaniaceae + Tropaeolaceae] [[Moringaceae + Caricaceae] [Setchellanthaceae

[Limnanthaceae [[Koeberliniaceae [Bataceae + Salvadoraceae]] [Emblingiaceae

[Pentadiplandraceae [Gyrostemonaceae Resedaceae Stixis and relatives] Tovariaceae

[Capparaceae [Cleomaceae + Brassicaceae]]]]]]]]

Berberidopsidales

Aextoxicaceae + Berberidopsidaceae

Santalales

Erythropalaceae Olacaceae [[Loranthaceae [Misodendraceae + Schoepfiaceae]]

[Opiliaceae + Santalaceae]] Balanophoraceae unplaced

Caryophyllales

[[Droseraceae [Nepenthaceae [Drosophyllaceae [Ancistrocladaceae +

Dioncophylleaceae]]]]] [[Frankeniaceae + Tamaricaceae] [Polygonaceae +

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

37

Plumbaginaceae]]] [Rhabdodendraceae [Simmondsiaceae [[Asteropeiaceae +

Physenaceae] [[Caryophyllaceae [Achatocarpaceae + Amaranthaceae]]

[Stegnospermataceae [Limeaceae [[Lophocarpaceae [Barbeuiaceae [Aizoaceae

[Sarcobataceae Phytolaccaceae Nyctaginaceae]]]] [Molluginaceae [Halophytaceae

Montiaceae Basellaceae Didiereaceae [Talinaceae [Portulacaceae [Anacampseros etc +

Cactaceae]]]]]]]]]]]]]

ASTERIDS

Cornales

[Cornaceae + Nyssaceae] [Hydrangeaceae + Loasaceae] Hydrostachyaceae [Curtisiaceae

+ Grubbiaceae]

Ericales

[Balsaminaceae [Marcgraviaceae + Tetrameristaceae]] [[Polemoniaceae +

Fouquieraceae] Lecythidaceae [[Sladeniaceae + Pentaphylacaceae] [Sapotaceae

[Ebenaceae [Maesaceae [Theophrastaceae [Myrsinaceae + Primulaceae]]]]]

[Mitrastemonaceae Theaceae [Symplocaceae [Styracaceae + Diapensiaceae]]

[[Sarraceniaceae [Roridulaceae + Actinidiaceae] [Clethraceae [Cyrillaceae +

Ericaceae]]]]]]

LAMIIDASTERID I

Unplaced

Metteniusaceae Oncothecaceae Icacinaceae and some unplaced ex-Icacinaceae

Garryales

Garryaceae + Eucommiaceae

Unplaced

Boraginaceae Vahliaceae

Gentianales

Rubiaceae [Gentianaceae [Loganiaceae [Gelsemiaceae + Apocynaceae]]]

Lamiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

38

Plocospermataceae [Oleaceae [Tetrachondraceae [Calceolariaceae

[Gesneriaceae Peltanthera [Pedaliaceae Carlemanniaceae Martyniaceae Stilbaceae

Plantaginaceae Linderniaceae Bignoniaceae Verbenaceae Lamiaceae Paulowniaceae

Schlegeliaceae Thomandersiaceae Phrymaceae Rehmannia Acanthaceae

Scrophulariaceae Orobanchaceae Byblidaceae Lentibulariaceae]]]]]

Solanales

[Montiniaceae [Sphenocleaceae + Hydroleaceae]] [Convolvulaceae + Solanaceae]

CAMPANULIDASTERID II

Aquifoliales

[[Cardiopteridaceae + Stemonuraceae] [Aquifoliaceae [Helwingiaceae +

Phyllonomaceae]]

Asterales

[Rousseaceae + Campanulaceae] Pentaphragmataceae [[Alseuosmiaceae [Phellinaceae

+ Argophyllaceae]] Stylidiaceae [Menyanthaceae [Goodeniaceae [Calyceraceae +

Asteraceae]]]]

Escalloniales

Escalloniaceae

Bruniales

[Bruniaceae + Columelliaceae]

Apiales

Pennantiaceae [Torricelliaceae [Griseliniaceae [Pittosporaceae [Araliaceae

[Myodocarpaceae + Apiaceae]]]]]

Paracryphiales

Paracryphiaceae

Dipsacales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

39

Adoxaceae [Diervillaceae [Caprifoliaceae [Linnaeaceae [Morinaceae [Dipsacaceae +

Valerianaceae]]]]]

LINKS TO ORDERS AND FAMILIES

[Back to Top]

MAIN GROUPINGS

Asterids commelinids core eudicots asterid 1 asterid

2 eudicots gymnosperms Magnoliophyta monocots N-fixing claderosids seed plants

ALPHABETICAL LISTING OF ALL ORDINAL NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | L | M | N | O | P | Q | R | S | T | U | V | W | X | Z

Abietales Acanthales Acerales Acorales Actinidiales Actinostrobales Adoxales Aescul

ales Agavales Aizoales AkanialesAlismatales Alliales Alseuosmiales Alstroemeriales

Altingiales Amaranthales Amaryllidales Amborellales AmbrosialesAmmiales Amomal

es Ancistrocladales Anisophylleales Annonales Anthobolales Apiales Apocynales Apo

nogetonalesAquifoliales Arales Araliales Aralidiales Araucariales Arecales Aristolochi

ales Athrotaxidales Asarales AsclepiadalesAsparagales Asphodelales Asparagales Ast

eliales Atriplicales Aucubales Austrobaileyales Avenales

Balanitales Balanopales Balanophorales Balsaminales Barbeyales Barclayales Batales

Begoniales BerberidalesBerberidopsidales Betulales Biebersteiniales Bignoniales Bix

ales Boraginales Brassicales Brexiales Bromeliales BrunialesBrunoniales Burmannial

es Burserales Butomales Buxales Byblidales

Cactales Callitrichales Calycanthales Calycerales Campanulales Campynematales Can

ellales Cannales CapparalesCaprifoliales Cardiopteridales Carduales Caricales Carle

manniales Caryophyllales Cassiales Casuarinales CelastralesCentrolepidales Cephalot

ales Cephalotaxales Ceratophyllales Cercidiphyllales Chenopodiales Chironiales Chlor

anthalesChrysobalanales Cinchonales Circaeasterales Cistales Citrales Cocosales Col

chicales Columelliales CombretalesCommelinales Connarales Convolvulales Coriarial

es Cornales Corylales Corynocarpales Crassulales CrossosomatalesCucurbitales Cunn

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

40

inghamiales Cunoniales Cupressales Cycadales Cyclanthales Cymodoceales Cynarales

CynomorialesCyperales Cyrillales Cytinales

Daphnales Daphniphyllales Dasypogonales Datiscales Degeneriales Desfontainiales D

iapensiales Didymelales DillenialesDioales Dioncophyllales Dioscoreales Diospyrales

Dipentodontales Dipsacales Droserales

Ebenales Echiales Elaeagnales Elaeocarpales Elatinales Elodeales Emmotales Empetr

ales Ephedrales Ericales EriocaulalesErythropalaless Escalloniales Eucommiales Euph

orbiales Eupomatiales Eupteleales Euryalaless

Fabales Fagales Falcatifoliales Ficales Flacourtiales Flagellariales Fouquieriales Franc

oales Frangulales

Galiales Garryales Geissolomatales Gentianales Geraniales Gesneriales Ginkgoales G

laucidiales Globulariales GnetalesGoodeniales Greyiales Griseliniales Grossulariales

Grubbiales Gunnerales Gyrocarpales Gyrostemonales

Haemodorales Haloragales Hamamelidales Hanguanales Heisteriales Helleborales He

lwingiales Himantandrales HippuridalesHomaliales Hortensiales Huales Huerteales

Hydatellales Hydnorales Hydrangeales Hydrastidales HydrocharitalesHydropeltidales

Hydrostachyales Hypericales Hypoxidales

Icacinales Illiciales Iridales Irvingales Iteales Ixiales

Jasminales Juglandales Julianiales Juncaginales Juncales

Lacistematales Lactoridales Lamiales Lardizabalales Laurales Lecythidales Ledocarpal

es Leitneriales LentibularialesLigustrales Liliales Limnanthales Linales Liriales Loasal

es Lobeliales Loganiales Lonicerales Loranthales LowialesLythrales

Magnoliales Malpighiales Malvales Marathrales Marcgraviales Mayacales Medusagy

nales Medusandrales MelanthialesMelastomatales Meliales Melianthales Meliosmal

es Menispermales Menyanthales Metteniusales MitrastemonalesMiyoshiales Moni

miales Moringales Musales Myricales Myristicales Myrothamnales Myrsinales Myrta

les

Najadales Nandinales Narcissales Nartheciales Nelumbonales Nepenthales Neuradal

es Nitrariales Nolanales NothofagalesNyctaginales Nymphaeales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

41

Ochnales Oenotherales Olacales Oleales Onagrales Oncothecales Opuntiales Orchid

ales Oxalidales

Saxifragales Pandales Pandanales Papaverales Paracryphiales Parasitaxales Paridales

Parnassiales PassifloralesPenaeales Pennantiales Pentaphragmatales Peridiscales P

etiveriales Petrosaviales Phellinales Philydrales PhyllanthalesPhysenales Phytolaccal

es Picramniales Pinales Pinguiculales Piperales Pittosporales Plantaginales Platanale

sPlumbaginales Poales Podocarpales Podophyllales Podostemales Polemoniales Pol

ygalales Polygonales PontederialesPortulacales Posidoniales Potamogetonales Primu

lales Proteales

Quercales Quillajales Quintiniales

Rafflesiales Ranunculales Rapateales Resedales Restionales Rhabdodendrales Rhamn

ales Rhinanthales RhizophoralesRhodorales Rhoipteleales Roridulales Rosales Rouss

eales Rubiales Ruppiales Rutales

Sabiales Salicales Salvadorales Samolales Samydales Sanguisorbales Santalales Sapin

dales Sapotales SarracenialesSaxegotheales Saxifragales Scheuchzeriales Sciadopitya

les Scleranthales Scrophulariales Scyphostegiales SedalesSilenales Simmondsiales S

milacales Solanales Sphenocleales Sphenostemonales Stangeriales Stellariales Stemo

nalesStilbales Stylidiales Styracales Surianales

Taccales Tamales Tamaricales Taxales Taxodiales Tecophilaeales Terebinthales Tern

stroemiales Theales TheligonalesThymelaeales Tiliales Tofieldiales Torricelliales Tov

ariales Tribelales Trilliales Trimeniales Triuridales TrochodendralesTropaeolales Turn

erales Typhales

Ulmales Urticales

Vacciniales Vahliales Vallisneriales Velloziales Veratrales Verbenales Viburnales Vinc

ales Violales Viscales VitalesVochysiales

Welwitschiales Winterales

Xanthorrhoeales Ximeniales Xyridales

Zamiales Zingiberales Zosterales Zygophyllales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

42

ALPHABETICAL LISTING OF ALL FAMILY NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | K | L | M | N | O | P | Q | R | S | T | U | V | W | X | Y

|Z

Back to Home

In some cases clicking on a family synonym will send you directly to the subfamily in

which the genus of the basionym is to be found

Abietaceae Abolbodaceae Abrophyllaceae Acalyphaceae Acanthaceae [Acanthochla

mydaceae] Acarnaceaee AceraceaeAchariaceae Achatocarpaceae Achradaceae Achy

ranthaceae Acmopylaceae Acoraceae Acristaceae ActaeaceaeActinidiaceae Actinota

ceae Amentotaxaceae Adenogrammaceae Adoxaceae Aegialitidaceae Aegicerataceae

Aegilopaceae Aeginetiaceaee Aegiphilaceae Aesculaceae Aextoxicaceae Agapanthac

eae Agavaceae AgdestidaceaeAgialidaceae Agrimoniaceae Agrostidaceae Ailanthace

ae Aitoniaceae Aizoaceae Akaniaceae Alangiaceae AlchemillaceaeAldrovandaceae A

lismataceae Alliaceae Allioniaceae Allophylaceae Aloaceae Alopecuraceae Alpiniacea

e AlseuosmiaceaeAlsinaceae Alsodeiaceae Alstroemeriaceae Altingiaceae Alzateacea

e Amaranthaceae Amaryllidaceae Amborellaceae Ambrosiaceae Amentotaxaceae A

mmanniaceae Ammiaceae Amomaceae Ampelidopsaceae Amygdalaceae Amyridacea

eAnacardiaceae Anagallidaceae Anarthriaceae Anchusaceae Ancistrocladaceae Andr

omedaceae AndropogonaceaeAndrostachyaceae Androsynaceae Anemarrhenaceae

Anemonaceae Angelicaceae Anisophylleaceae AnnonaceaeAnomochloaceae Anopter

aceae Anrederaceae Anthemidaceae Anthericaceae Anthobolaceae Antidesmataceae

Antirrhinaceae Antoniaceae Aparinaceae Aphanopetalaceae Aphloiaceae Aphyllanth

aceae Apiaceae ApocynaceaeApodanthaceae Aponogetonaceae Aporusaceae Apose

ridaceae Apostasiaceae Aptandraceae AquifoliaceaeAquilariaceae Aquilegiaceae A

raceae Aragoaceae Araliaceae Aralidiaceaee Araucariaceae Arbutaceae Arceuthidace

aeArceuthobiaceae Arctostaphylaceae Arctotidaceae Ardisiaceae Arecaceae Argoph

yllaceae Arisaraceae AristolochiaceaeAristoteliaceae Arjonaceae Armeriaceae Artem

isiaceae Arthrotaxidaceae Artocarpaceae Arundinaceae ArundinellaceaeAsaraceae A

sclepiadaceae Ascyraceaee Asiraceae Asparagaceae Asperulaceae Asphodelaceae As

pidistraceaeeAsteliaceae Asteraceae Asteranthaceae Asteropeiaceae Astragalaceae

Astrocarpaceae AthanasiaceaeAtherospermataceae Atriplicaceae Atropaceae Aucuba

ceae Aurantiaceae Austrobaileyaceae Austrotaxaceae AvenaceaeAverrhoaceae Avet

raceae Avicenniaceae Azaleaceae Azimaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

43

Balanitaceae Balanopaceae Balanophoraceae Balsameaceae Balsaminaceae Bambusa

ceae Barbaceniaceae BarbeuiaceaeBarbeyaceae Barclayaceae Barringtoniaceae Base

llaceae Bataceae Baueraceae Bauhiniaceae Baxteriaceae BegoniaceaeBehniaceae B

elangeraceae Belloniaceae Bembiciaceae Berberidaceae Berberidopsidaceae Berryac

eaee BersamaceaeBertyaceae Berzeliaceae Besleriaceae Betaceae Betulaceae Biebe

rsteiniaceae Bifariaceae Bignoniaceae BischofiaceaeBixaceae Blakeaceae Blandfordia

ceae Blattiaceae Blepharocaryaceae Blitaceae Blyxaceae Boerlagellaceae Bolivaracea

eBombacaceae Bonnetiaceae Bontiaceae Boopidaceae Boraginaceae Borassaceae B

oroniaceae BoryaceaeBotryodendraceae Bougainvilleaceae Boweniaceae Brassicacea

e Brachycaulaceae Bretschneideraceae BrexiaceaeBromeliaceae Brownlowiaceae Br

unelliaceae Bruniaceae Brunoniaceae Brunsvigiaceae Bryoniaceae BuchneraceaeBuci

daceae Buddlejaceae Buglossaceae Bulbocodiaceae Bumeliaceae Bupleuraceae Burc

hardiaceae BurmanniaceaeBurseraceae Butneriaceae Butomaceae Buxaceae Byblida

ceae Byttneriaceae

Cabombaceae Cacaoaceae Cactaceae Caesalpiniaceae Caladiaceae Calamaceae Calce

olariaceae CalectasiaceaeCalendulaceae Callaceae Callicomaceae Calligonaceae Calli

traceae Callitrichaceae Calochortaceae CalophyllaceaeCalthaceae Calycanthaceae Ca

lyceraceae Cambogiaceae Camelliaceae Campanulaceae CampynemataceaeCanacom

yricaceae Canellaceae Cannabaceae Cannaceae Canopodaceae Canotiaceae Cansjera

ceae CapparaceaeCaprariaceae Caprifoliaceae Cardiopteridaceae Carduaceae Carica

ceae Carissaceae Carlemanniaceae CarpinaceaeCarpodetaceae Cartonemataceae Ca

ryocaraceae Caryophyllaceae Casearieaceae Cassiaceae CassipoureaceaeCassythacea

e Castaneaceae Castelaceae Casuarinaceae Catesbaeaceae Cathedraceae Cecropiace

ae CedraceaeCedrelaceae Celastraceae Celosiaceae Celtidaceae Centaureaceae Cen

trolepidaceae Centroplacaceae CepaceaeCephalanthaceae Cephalotaceae Cephalota

xaceae Cerastiaceae Ceratoniaceae Ceratophyllaceae CerberaceaeCercidiphyllaceae

Cercocarpaceae Cercodiaceae Cereaceae Cerinthaceae Ceroxylaceae Cestraceae Cev

alliaceaeChailletiaceae Chamaedoraceae Chamaemoraceae Chamelauciaceae Chaun

ochitonaceae Chelidoniaceae ChelonaceaeChenopodiaceae Chimonanthaceae Chingi

thamnaceae Chionographidaceae Chiranthodendraceae ChironiaceaeChlaenaceae Ch

loanthaceae Chloranthaceae Chloridaceae Chlorogalaceae Chrysobalanaceae Cicerac

eae CichoriaceaeCimicifugaceae Cinchoniaceae Circaeaceae Circaeasteraceae Cissac

eae Cistaceae Citraceae Clematidaceae CleomaceaeClethraceae Clusiaceae Cneorac

eae Cnestidaceae Cnicaceae Cobaeaceae Cochlospermaceae Cocosaceae Coffeaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

44

Colchicaceae Coleogynaceae Columelliaceae Combretaceae Commelinaceae Comocl

adiaceae Compositae CompsoaceaeConantheraceae Connaraceae Conostylidaceae C

onvallariaceae Convolvulaceae Cordiaceae CoreopsidaceaeCoriandraceae Coriariacea

e Coridaceae Corispermaceae Cornaceae Corokiaceae Coronillaceae Correaceae Cor

rigiolaceaeCorsiaceae Corydalaceae Corylaceae Corynocarpaceae Coryphaceae Cost

aceae Cotyledonaceae CoulaceaeCoutariaceae Coutoubiaceae Crassulaceae Crescen

tiaceae Cressaceae Crinaceae Crocaceae CroomiaceaeCrossosomataceae Crotonacea

e Cruciferae Cryptaceae Crypteroniaceae Cryptocorynaceae CryptomeriaceaeCtenol

ophonaceae Cucurbitaceae Cunninghamiaceae Cunoniaceae Cupressaceae Curcumac

eae Curtisiaceae CuscutaceaeCyananthaceae Cyanastraceae Cynanchaceae Cyanella

ceae Cycadaceae Cyclanthaceae CyclantheraceaeCyclocheilaceae Cydoniaceae Cymo

doceaceae Cynaraceae Cynocrambaceae Cynomoriaceae Cyperaceae CyphiaceaeCyp

hocarpaceae Cypripediaceae Cyrillaceae Cyrtandraceae Cyrtanthaceae Cytinaceae

Dacrycarpaceae Dactylanthaceae Damasoniaceae Daphnaceae Daphniphyllaceae Das

ypogonaceae DatiscaceaeDaturaceae Daucaceae Davidiaceae Davidsoniaceae Decais

neaceae Deeringiaceae Degeneriaceae DelphiniaceaeDendrophthoaceae Columelliac

eae Detariaceae Dialypetalanthaceae Dianellaceae Dianthaceae DiapensiaceaeDicha

petalaceae Dichondraceae Diclidantheraceae Dicrastylidiaceae Dictamnaceae Didiere

aceae DidymelaceaeDidymocarpaceae Diegodendraceae Diervillaceae Digitalidaceae

e Dilatridaceae Dilleniaceae Dioaceae DionaeaceaeDioncophyllaceae Dioscoreaceae

Diosmaceae Diospyraceae Dipentodontaceae Diphylleiaceae Diplolaenaceae Dipsaca

ceaeDipterocarpaceae Dirachmaceae Disanthaceae Diselmaceae Dodonaeaceae Do

mbeyaceae Donatiaceae DorsteniaceaeDortmannaceae Doryanthaceae Drabaceae D

racaenaceae Dracontiaceae Drimyidaceae Droseraceae DrosophyllaceaeDryadaceae

Duabangaceae Duckeodendraceae Dulongiaceae Durantaceae Durionaceae Dysphani

aceae

Ebenaceae Eccremidaceae Ecdeiocoleaceae Echiaceae Echinopaceae Ehretiaceae Ela

eagnaceae ElaeocarpaceaeElegiaceae Elatinaceae Ellisiaceae Ellisiophyllaceae Elodea

ceae Elytranthaceae Embeliaceae Emblingiaceae EmmotaceaeEmpetraceae Encepha

lartaceae Engelhardtiaceae Enhalaceae Epacridaceae Ephedraceae Ephemeranthacea

eEpilobiaceae Epimediaceae Eragrostidaceae Eremolepidaceae Eremosynaceae Erica

ceae Erinaceae EriocaulaceaeEriogonaceae Eriospermaceae Erodiaceae Erycibaceae

Eryngiaceae Erysimaceae Erythroniaceae ErythropalaceaeErythrospermaceae Erythro

xylaceae Escalloniaceae Eschscholziaceae Eucomidaceae Eucommiaceae Eucryphiace

aeEuonymaceae Eupatoriaceae Euphorbiaceae Euphrasiaceae Euphroniaceae Eupo

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

45

matiaceae Eupteleaceae EuryalaceaeEustrephaceae Euthemidaceae Eutocaceae Exb

ucklandiaceae Exocarpaceae

Fabaceae Fagaceae Falcatifoliaceae Ferulaceae Festucaceae Ficaceae Ficoidaceae Fit

zroyaceae FlacourtiaceaeFlagellariaceae Flindersiaceae Foetidiaceae Forestieraceae

Fothergillaceae Fouquieriaceae Fragariaceae FrancoaceaeFrangulaceae Frankeniacea

e Fraxinaceae Fraxinellaceae Freycinetiaceae Fritillariaceae Fuchsiaceae Fumariacea

eFunkiaceae

Gaiadendraceae Galacaceae Galanthaceae Galaxiaceae Galedupaceae Galeniaceae G

aliaceae Garciniaceae GardeniaceaeGardneriaceae Garryaceae Geissolomataceae Ge

itonoplesiaceae Gelsemiaceae Geniostomaceae GentianaceaeGeonomataceae Geosir

idaceae Geraniaceae Gerrardinaceae Gesneriaceae Gethyllidaceae Gilliesiaceae Gina

lloaceaeGinkgoaceae Gisekiaceae Gladiolaceae Glaucidiaceae Glechomaceae Glinace

ae Globulariaceae Gnaphaliaceae GnetaceaeGoetzeaceae Gomortegaceae Gomphiac

eae Gomphrenaceae Gonystylaceae Goodeniaceae Gordoniaceae GouaniaceaeGoup

iaceae Gramineae Gratiolaceae Grewiaceae Greyiaceae Grielaceae Griseliniaceae Gr

onoviaceae GrossulariaceaeGrubbiaceae Guaiacanaceae Guamatelaceae Guettardac

eae Gunneraceae Gustaviaceae Guttiferae GyrocarpaceaeGyrostemonaceae

Hachetteaceae Haemanthaceae Haemodoraceae Halesiaceae Hallieraceae Halocarpa

ceae Halophilaceae HalophytaceaeHaloragaceae Hamamelidaceae Hanguanaceae H

aptanthaceae Harmandaceae Hebenstretiaceae HectorellaceaeHederaceae Hedyosm

aceae Hedyotidaceae Hedysaraceae Heisteriaceae Heleniaceae Heliamphoraceae H

elianthaceaeHelianthemaceae Helichrysaceae Heliconiaceae Helicteraceae Heliotropi

aceae Helleboraceae Heloniadaceae HelosidaceaeHelwingiaceae Hemerocallidaceae

Hemimeridaceae Hemiphylacaceae Henriqueziaceae Henslowiaceae Hermanniaceae

Hernandiaceae Herniariaceae Herreriaceae Hesperocallidaceae Heterantheraceae He

teropyxidaceae HeterostylaceaeHibbertiaceae Hibiscaceae Hilleriaceae Himantandra

ceae Hippocastanaceae Hippocrateaceae HippomanaceaeHippophaeaceae Hippurida

ceae Hirtellaceae Holacanthaceae Homaliaceae Hopkinsiaceae Hoplestigmataceae H

ordeaceaeHornschuchiaceae Hortensiaceae Hortoniaceae Hostaceae Houstoniaceae

Huaceae Huerteaceae HugoniaceaeHumbertiaceae Humiriaceae Hyacinthaceae Hyd

atellaceae Hydnoraceae Hydrangeaceae Hydrastidaceae HydrillaceaeHydrocharitacea

e Hydrocotylaceae Hydrogetonaceae Hydroleaceae Hydropeltidaceae Hydrophylacac

eae HydrophyllaceaeHydrostachyaceae Hymenocardiaceae Hypecoaceae Hyperanth

eraceae Hypericaceae Hypopityaceae HypoxidaceaeHypseocharitaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

46

Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta

ceae ImperatoriaceaeInocarpaceae Inulaceae Ionidiaceae Iriarteaceae Iridaceae Irvi

ngiaceae Isnardiaceae Isophysidaceae IteaceaeStrasburgeriaceae Ixiaceae Ixioliriace

ae Ixonanthaceae

Jalapaceae Jamboliferaceae Japonoliriaceae Jasionaceae Jasminiaceae Johnsoniaceae

Joinvilleaceae JuglandaceaeJulianiaceae Juncaceae Juncaginaceae Juniperaceae Jus

siaeaceae Justiciaceae

Kadsuraceae Kaliphoraceae Kalmiaceae Kaniaceae Kiggelariaceae Kingdoniaceae Kin

giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae

Krameriaceae

Labiatae Lacandoniaceae Lachenaliaceae Lacistemataceae Lactoridaceae Lactucaceae

Lagerstroemiaceaee LamiaceaeLampsanaceae Lanariaceae Langsdorffiaceae Lantan

aceae Lapageriaceae Lardizabalaceae Lasiopetalaceae LauraceaeLawsoniaceae Laxm

anniaceae Lecythidaceae Ledaceae Ledocarpaceae Leeaceae Leguminosae Leitneriac

eae LemnaceaeLennoaceae Lentibulariaceae Lentiscaceae Leoniaceae Leonticaceae

Lepidobotryaceae LepidocarpaceaeLepidocaryaceae Lepidocerataceae Leptaulaceae

Leptospermaceae Lepturaceae Lepuropetalaceae LeuctenbergiaceaeLibocedraceae

Licaniaceae Ligustraceae Lilaeaceae Liliaceae Limeaceae Limnanthaceae Limnocharit

aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace

ae Linderniaceae Lindleyaceae Linnaeaceae LippayaceaeLiriaceae Liriodendraceae Li

ssocarpaceae Littorellaceae Loasaceae Lobeliaceaee Loganiaceae LomandraceaeLoni

ceraceae Lopeziaceae Lophiocarpaceae Lophiolaceae Lophiraceae Lophophytaceae L

ophopyxidaceae LoranthaceaeLotaceae Lowiaceae Lupulaceaee Luxemburgiaceae Lu

zuriagaceae Lychnidaceae Lyciaceae LyginiaceaeLygodisodeaceae Lysimachiaceae Ly

thraceae

Macarisiaceae Mackinlayaceae Madiaceae Maesaceae Magnoliaceae Malaceae Male

sherbiaceae MalortieaceaeMalpighiaceae Malvaceae Mangiaceae Manicariaceae Ma

paniaceae Marantaceae Marathraceae MarcgraviaceaeMartyniaceae Mastixiaceae

Matricariaceae Maundiaceae Mayacaceae Medeolaceae Medusagynaceae Peridiscac

eaeMelaleucaceae Melampyraceae Melanophyllaceae Melanthiaceae Melastomatac

eae Meliaceae Melianthaceae MelicaceaeMeliosmaceae Melittidaceae Melochiacea

e Memecylaceae Mendonciaceae Menispermaceae Menthaceae MenyanthaceaeMe

nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

47

Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

Nypaceae Nyssaceae

Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

ceae Papilionaceae Pappophoraceae PapyraceaeParacryphiaceae Parasitaxaceae Par

ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

48

hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

49

cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

50

ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 37: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

37

Plumbaginaceae]]] [Rhabdodendraceae [Simmondsiaceae [[Asteropeiaceae +

Physenaceae] [[Caryophyllaceae [Achatocarpaceae + Amaranthaceae]]

[Stegnospermataceae [Limeaceae [[Lophocarpaceae [Barbeuiaceae [Aizoaceae

[Sarcobataceae Phytolaccaceae Nyctaginaceae]]]] [Molluginaceae [Halophytaceae

Montiaceae Basellaceae Didiereaceae [Talinaceae [Portulacaceae [Anacampseros etc +

Cactaceae]]]]]]]]]]]]]

ASTERIDS

Cornales

[Cornaceae + Nyssaceae] [Hydrangeaceae + Loasaceae] Hydrostachyaceae [Curtisiaceae

+ Grubbiaceae]

Ericales

[Balsaminaceae [Marcgraviaceae + Tetrameristaceae]] [[Polemoniaceae +

Fouquieraceae] Lecythidaceae [[Sladeniaceae + Pentaphylacaceae] [Sapotaceae

[Ebenaceae [Maesaceae [Theophrastaceae [Myrsinaceae + Primulaceae]]]]]

[Mitrastemonaceae Theaceae [Symplocaceae [Styracaceae + Diapensiaceae]]

[[Sarraceniaceae [Roridulaceae + Actinidiaceae] [Clethraceae [Cyrillaceae +

Ericaceae]]]]]]

LAMIIDASTERID I

Unplaced

Metteniusaceae Oncothecaceae Icacinaceae and some unplaced ex-Icacinaceae

Garryales

Garryaceae + Eucommiaceae

Unplaced

Boraginaceae Vahliaceae

Gentianales

Rubiaceae [Gentianaceae [Loganiaceae [Gelsemiaceae + Apocynaceae]]]

Lamiales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

38

Plocospermataceae [Oleaceae [Tetrachondraceae [Calceolariaceae

[Gesneriaceae Peltanthera [Pedaliaceae Carlemanniaceae Martyniaceae Stilbaceae

Plantaginaceae Linderniaceae Bignoniaceae Verbenaceae Lamiaceae Paulowniaceae

Schlegeliaceae Thomandersiaceae Phrymaceae Rehmannia Acanthaceae

Scrophulariaceae Orobanchaceae Byblidaceae Lentibulariaceae]]]]]

Solanales

[Montiniaceae [Sphenocleaceae + Hydroleaceae]] [Convolvulaceae + Solanaceae]

CAMPANULIDASTERID II

Aquifoliales

[[Cardiopteridaceae + Stemonuraceae] [Aquifoliaceae [Helwingiaceae +

Phyllonomaceae]]

Asterales

[Rousseaceae + Campanulaceae] Pentaphragmataceae [[Alseuosmiaceae [Phellinaceae

+ Argophyllaceae]] Stylidiaceae [Menyanthaceae [Goodeniaceae [Calyceraceae +

Asteraceae]]]]

Escalloniales

Escalloniaceae

Bruniales

[Bruniaceae + Columelliaceae]

Apiales

Pennantiaceae [Torricelliaceae [Griseliniaceae [Pittosporaceae [Araliaceae

[Myodocarpaceae + Apiaceae]]]]]

Paracryphiales

Paracryphiaceae

Dipsacales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

39

Adoxaceae [Diervillaceae [Caprifoliaceae [Linnaeaceae [Morinaceae [Dipsacaceae +

Valerianaceae]]]]]

LINKS TO ORDERS AND FAMILIES

[Back to Top]

MAIN GROUPINGS

Asterids commelinids core eudicots asterid 1 asterid

2 eudicots gymnosperms Magnoliophyta monocots N-fixing claderosids seed plants

ALPHABETICAL LISTING OF ALL ORDINAL NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | L | M | N | O | P | Q | R | S | T | U | V | W | X | Z

Abietales Acanthales Acerales Acorales Actinidiales Actinostrobales Adoxales Aescul

ales Agavales Aizoales AkanialesAlismatales Alliales Alseuosmiales Alstroemeriales

Altingiales Amaranthales Amaryllidales Amborellales AmbrosialesAmmiales Amomal

es Ancistrocladales Anisophylleales Annonales Anthobolales Apiales Apocynales Apo

nogetonalesAquifoliales Arales Araliales Aralidiales Araucariales Arecales Aristolochi

ales Athrotaxidales Asarales AsclepiadalesAsparagales Asphodelales Asparagales Ast

eliales Atriplicales Aucubales Austrobaileyales Avenales

Balanitales Balanopales Balanophorales Balsaminales Barbeyales Barclayales Batales

Begoniales BerberidalesBerberidopsidales Betulales Biebersteiniales Bignoniales Bix

ales Boraginales Brassicales Brexiales Bromeliales BrunialesBrunoniales Burmannial

es Burserales Butomales Buxales Byblidales

Cactales Callitrichales Calycanthales Calycerales Campanulales Campynematales Can

ellales Cannales CapparalesCaprifoliales Cardiopteridales Carduales Caricales Carle

manniales Caryophyllales Cassiales Casuarinales CelastralesCentrolepidales Cephalot

ales Cephalotaxales Ceratophyllales Cercidiphyllales Chenopodiales Chironiales Chlor

anthalesChrysobalanales Cinchonales Circaeasterales Cistales Citrales Cocosales Col

chicales Columelliales CombretalesCommelinales Connarales Convolvulales Coriarial

es Cornales Corylales Corynocarpales Crassulales CrossosomatalesCucurbitales Cunn

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

40

inghamiales Cunoniales Cupressales Cycadales Cyclanthales Cymodoceales Cynarales

CynomorialesCyperales Cyrillales Cytinales

Daphnales Daphniphyllales Dasypogonales Datiscales Degeneriales Desfontainiales D

iapensiales Didymelales DillenialesDioales Dioncophyllales Dioscoreales Diospyrales

Dipentodontales Dipsacales Droserales

Ebenales Echiales Elaeagnales Elaeocarpales Elatinales Elodeales Emmotales Empetr

ales Ephedrales Ericales EriocaulalesErythropalaless Escalloniales Eucommiales Euph

orbiales Eupomatiales Eupteleales Euryalaless

Fabales Fagales Falcatifoliales Ficales Flacourtiales Flagellariales Fouquieriales Franc

oales Frangulales

Galiales Garryales Geissolomatales Gentianales Geraniales Gesneriales Ginkgoales G

laucidiales Globulariales GnetalesGoodeniales Greyiales Griseliniales Grossulariales

Grubbiales Gunnerales Gyrocarpales Gyrostemonales

Haemodorales Haloragales Hamamelidales Hanguanales Heisteriales Helleborales He

lwingiales Himantandrales HippuridalesHomaliales Hortensiales Huales Huerteales

Hydatellales Hydnorales Hydrangeales Hydrastidales HydrocharitalesHydropeltidales

Hydrostachyales Hypericales Hypoxidales

Icacinales Illiciales Iridales Irvingales Iteales Ixiales

Jasminales Juglandales Julianiales Juncaginales Juncales

Lacistematales Lactoridales Lamiales Lardizabalales Laurales Lecythidales Ledocarpal

es Leitneriales LentibularialesLigustrales Liliales Limnanthales Linales Liriales Loasal

es Lobeliales Loganiales Lonicerales Loranthales LowialesLythrales

Magnoliales Malpighiales Malvales Marathrales Marcgraviales Mayacales Medusagy

nales Medusandrales MelanthialesMelastomatales Meliales Melianthales Meliosmal

es Menispermales Menyanthales Metteniusales MitrastemonalesMiyoshiales Moni

miales Moringales Musales Myricales Myristicales Myrothamnales Myrsinales Myrta

les

Najadales Nandinales Narcissales Nartheciales Nelumbonales Nepenthales Neuradal

es Nitrariales Nolanales NothofagalesNyctaginales Nymphaeales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Ochnales Oenotherales Olacales Oleales Onagrales Oncothecales Opuntiales Orchid

ales Oxalidales

Saxifragales Pandales Pandanales Papaverales Paracryphiales Parasitaxales Paridales

Parnassiales PassifloralesPenaeales Pennantiales Pentaphragmatales Peridiscales P

etiveriales Petrosaviales Phellinales Philydrales PhyllanthalesPhysenales Phytolaccal

es Picramniales Pinales Pinguiculales Piperales Pittosporales Plantaginales Platanale

sPlumbaginales Poales Podocarpales Podophyllales Podostemales Polemoniales Pol

ygalales Polygonales PontederialesPortulacales Posidoniales Potamogetonales Primu

lales Proteales

Quercales Quillajales Quintiniales

Rafflesiales Ranunculales Rapateales Resedales Restionales Rhabdodendrales Rhamn

ales Rhinanthales RhizophoralesRhodorales Rhoipteleales Roridulales Rosales Rouss

eales Rubiales Ruppiales Rutales

Sabiales Salicales Salvadorales Samolales Samydales Sanguisorbales Santalales Sapin

dales Sapotales SarracenialesSaxegotheales Saxifragales Scheuchzeriales Sciadopitya

les Scleranthales Scrophulariales Scyphostegiales SedalesSilenales Simmondsiales S

milacales Solanales Sphenocleales Sphenostemonales Stangeriales Stellariales Stemo

nalesStilbales Stylidiales Styracales Surianales

Taccales Tamales Tamaricales Taxales Taxodiales Tecophilaeales Terebinthales Tern

stroemiales Theales TheligonalesThymelaeales Tiliales Tofieldiales Torricelliales Tov

ariales Tribelales Trilliales Trimeniales Triuridales TrochodendralesTropaeolales Turn

erales Typhales

Ulmales Urticales

Vacciniales Vahliales Vallisneriales Velloziales Veratrales Verbenales Viburnales Vinc

ales Violales Viscales VitalesVochysiales

Welwitschiales Winterales

Xanthorrhoeales Ximeniales Xyridales

Zamiales Zingiberales Zosterales Zygophyllales

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ALPHABETICAL LISTING OF ALL FAMILY NAMES OF SEED PLANTS WITH LINKS

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In some cases clicking on a family synonym will send you directly to the subfamily in

which the genus of the basionym is to be found

Abietaceae Abolbodaceae Abrophyllaceae Acalyphaceae Acanthaceae [Acanthochla

mydaceae] Acarnaceaee AceraceaeAchariaceae Achatocarpaceae Achradaceae Achy

ranthaceae Acmopylaceae Acoraceae Acristaceae ActaeaceaeActinidiaceae Actinota

ceae Amentotaxaceae Adenogrammaceae Adoxaceae Aegialitidaceae Aegicerataceae

Aegilopaceae Aeginetiaceaee Aegiphilaceae Aesculaceae Aextoxicaceae Agapanthac

eae Agavaceae AgdestidaceaeAgialidaceae Agrimoniaceae Agrostidaceae Ailanthace

ae Aitoniaceae Aizoaceae Akaniaceae Alangiaceae AlchemillaceaeAldrovandaceae A

lismataceae Alliaceae Allioniaceae Allophylaceae Aloaceae Alopecuraceae Alpiniacea

e AlseuosmiaceaeAlsinaceae Alsodeiaceae Alstroemeriaceae Altingiaceae Alzateacea

e Amaranthaceae Amaryllidaceae Amborellaceae Ambrosiaceae Amentotaxaceae A

mmanniaceae Ammiaceae Amomaceae Ampelidopsaceae Amygdalaceae Amyridacea

eAnacardiaceae Anagallidaceae Anarthriaceae Anchusaceae Ancistrocladaceae Andr

omedaceae AndropogonaceaeAndrostachyaceae Androsynaceae Anemarrhenaceae

Anemonaceae Angelicaceae Anisophylleaceae AnnonaceaeAnomochloaceae Anopter

aceae Anrederaceae Anthemidaceae Anthericaceae Anthobolaceae Antidesmataceae

Antirrhinaceae Antoniaceae Aparinaceae Aphanopetalaceae Aphloiaceae Aphyllanth

aceae Apiaceae ApocynaceaeApodanthaceae Aponogetonaceae Aporusaceae Apose

ridaceae Apostasiaceae Aptandraceae AquifoliaceaeAquilariaceae Aquilegiaceae A

raceae Aragoaceae Araliaceae Aralidiaceaee Araucariaceae Arbutaceae Arceuthidace

aeArceuthobiaceae Arctostaphylaceae Arctotidaceae Ardisiaceae Arecaceae Argoph

yllaceae Arisaraceae AristolochiaceaeAristoteliaceae Arjonaceae Armeriaceae Artem

isiaceae Arthrotaxidaceae Artocarpaceae Arundinaceae ArundinellaceaeAsaraceae A

sclepiadaceae Ascyraceaee Asiraceae Asparagaceae Asperulaceae Asphodelaceae As

pidistraceaeeAsteliaceae Asteraceae Asteranthaceae Asteropeiaceae Astragalaceae

Astrocarpaceae AthanasiaceaeAtherospermataceae Atriplicaceae Atropaceae Aucuba

ceae Aurantiaceae Austrobaileyaceae Austrotaxaceae AvenaceaeAverrhoaceae Avet

raceae Avicenniaceae Azaleaceae Azimaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Balanitaceae Balanopaceae Balanophoraceae Balsameaceae Balsaminaceae Bambusa

ceae Barbaceniaceae BarbeuiaceaeBarbeyaceae Barclayaceae Barringtoniaceae Base

llaceae Bataceae Baueraceae Bauhiniaceae Baxteriaceae BegoniaceaeBehniaceae B

elangeraceae Belloniaceae Bembiciaceae Berberidaceae Berberidopsidaceae Berryac

eaee BersamaceaeBertyaceae Berzeliaceae Besleriaceae Betaceae Betulaceae Biebe

rsteiniaceae Bifariaceae Bignoniaceae BischofiaceaeBixaceae Blakeaceae Blandfordia

ceae Blattiaceae Blepharocaryaceae Blitaceae Blyxaceae Boerlagellaceae Bolivaracea

eBombacaceae Bonnetiaceae Bontiaceae Boopidaceae Boraginaceae Borassaceae B

oroniaceae BoryaceaeBotryodendraceae Bougainvilleaceae Boweniaceae Brassicacea

e Brachycaulaceae Bretschneideraceae BrexiaceaeBromeliaceae Brownlowiaceae Br

unelliaceae Bruniaceae Brunoniaceae Brunsvigiaceae Bryoniaceae BuchneraceaeBuci

daceae Buddlejaceae Buglossaceae Bulbocodiaceae Bumeliaceae Bupleuraceae Burc

hardiaceae BurmanniaceaeBurseraceae Butneriaceae Butomaceae Buxaceae Byblida

ceae Byttneriaceae

Cabombaceae Cacaoaceae Cactaceae Caesalpiniaceae Caladiaceae Calamaceae Calce

olariaceae CalectasiaceaeCalendulaceae Callaceae Callicomaceae Calligonaceae Calli

traceae Callitrichaceae Calochortaceae CalophyllaceaeCalthaceae Calycanthaceae Ca

lyceraceae Cambogiaceae Camelliaceae Campanulaceae CampynemataceaeCanacom

yricaceae Canellaceae Cannabaceae Cannaceae Canopodaceae Canotiaceae Cansjera

ceae CapparaceaeCaprariaceae Caprifoliaceae Cardiopteridaceae Carduaceae Carica

ceae Carissaceae Carlemanniaceae CarpinaceaeCarpodetaceae Cartonemataceae Ca

ryocaraceae Caryophyllaceae Casearieaceae Cassiaceae CassipoureaceaeCassythacea

e Castaneaceae Castelaceae Casuarinaceae Catesbaeaceae Cathedraceae Cecropiace

ae CedraceaeCedrelaceae Celastraceae Celosiaceae Celtidaceae Centaureaceae Cen

trolepidaceae Centroplacaceae CepaceaeCephalanthaceae Cephalotaceae Cephalota

xaceae Cerastiaceae Ceratoniaceae Ceratophyllaceae CerberaceaeCercidiphyllaceae

Cercocarpaceae Cercodiaceae Cereaceae Cerinthaceae Ceroxylaceae Cestraceae Cev

alliaceaeChailletiaceae Chamaedoraceae Chamaemoraceae Chamelauciaceae Chaun

ochitonaceae Chelidoniaceae ChelonaceaeChenopodiaceae Chimonanthaceae Chingi

thamnaceae Chionographidaceae Chiranthodendraceae ChironiaceaeChlaenaceae Ch

loanthaceae Chloranthaceae Chloridaceae Chlorogalaceae Chrysobalanaceae Cicerac

eae CichoriaceaeCimicifugaceae Cinchoniaceae Circaeaceae Circaeasteraceae Cissac

eae Cistaceae Citraceae Clematidaceae CleomaceaeClethraceae Clusiaceae Cneorac

eae Cnestidaceae Cnicaceae Cobaeaceae Cochlospermaceae Cocosaceae Coffeaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Colchicaceae Coleogynaceae Columelliaceae Combretaceae Commelinaceae Comocl

adiaceae Compositae CompsoaceaeConantheraceae Connaraceae Conostylidaceae C

onvallariaceae Convolvulaceae Cordiaceae CoreopsidaceaeCoriandraceae Coriariacea

e Coridaceae Corispermaceae Cornaceae Corokiaceae Coronillaceae Correaceae Cor

rigiolaceaeCorsiaceae Corydalaceae Corylaceae Corynocarpaceae Coryphaceae Cost

aceae Cotyledonaceae CoulaceaeCoutariaceae Coutoubiaceae Crassulaceae Crescen

tiaceae Cressaceae Crinaceae Crocaceae CroomiaceaeCrossosomataceae Crotonacea

e Cruciferae Cryptaceae Crypteroniaceae Cryptocorynaceae CryptomeriaceaeCtenol

ophonaceae Cucurbitaceae Cunninghamiaceae Cunoniaceae Cupressaceae Curcumac

eae Curtisiaceae CuscutaceaeCyananthaceae Cyanastraceae Cynanchaceae Cyanella

ceae Cycadaceae Cyclanthaceae CyclantheraceaeCyclocheilaceae Cydoniaceae Cymo

doceaceae Cynaraceae Cynocrambaceae Cynomoriaceae Cyperaceae CyphiaceaeCyp

hocarpaceae Cypripediaceae Cyrillaceae Cyrtandraceae Cyrtanthaceae Cytinaceae

Dacrycarpaceae Dactylanthaceae Damasoniaceae Daphnaceae Daphniphyllaceae Das

ypogonaceae DatiscaceaeDaturaceae Daucaceae Davidiaceae Davidsoniaceae Decais

neaceae Deeringiaceae Degeneriaceae DelphiniaceaeDendrophthoaceae Columelliac

eae Detariaceae Dialypetalanthaceae Dianellaceae Dianthaceae DiapensiaceaeDicha

petalaceae Dichondraceae Diclidantheraceae Dicrastylidiaceae Dictamnaceae Didiere

aceae DidymelaceaeDidymocarpaceae Diegodendraceae Diervillaceae Digitalidaceae

e Dilatridaceae Dilleniaceae Dioaceae DionaeaceaeDioncophyllaceae Dioscoreaceae

Diosmaceae Diospyraceae Dipentodontaceae Diphylleiaceae Diplolaenaceae Dipsaca

ceaeDipterocarpaceae Dirachmaceae Disanthaceae Diselmaceae Dodonaeaceae Do

mbeyaceae Donatiaceae DorsteniaceaeDortmannaceae Doryanthaceae Drabaceae D

racaenaceae Dracontiaceae Drimyidaceae Droseraceae DrosophyllaceaeDryadaceae

Duabangaceae Duckeodendraceae Dulongiaceae Durantaceae Durionaceae Dysphani

aceae

Ebenaceae Eccremidaceae Ecdeiocoleaceae Echiaceae Echinopaceae Ehretiaceae Ela

eagnaceae ElaeocarpaceaeElegiaceae Elatinaceae Ellisiaceae Ellisiophyllaceae Elodea

ceae Elytranthaceae Embeliaceae Emblingiaceae EmmotaceaeEmpetraceae Encepha

lartaceae Engelhardtiaceae Enhalaceae Epacridaceae Ephedraceae Ephemeranthacea

eEpilobiaceae Epimediaceae Eragrostidaceae Eremolepidaceae Eremosynaceae Erica

ceae Erinaceae EriocaulaceaeEriogonaceae Eriospermaceae Erodiaceae Erycibaceae

Eryngiaceae Erysimaceae Erythroniaceae ErythropalaceaeErythrospermaceae Erythro

xylaceae Escalloniaceae Eschscholziaceae Eucomidaceae Eucommiaceae Eucryphiace

aeEuonymaceae Eupatoriaceae Euphorbiaceae Euphrasiaceae Euphroniaceae Eupo

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matiaceae Eupteleaceae EuryalaceaeEustrephaceae Euthemidaceae Eutocaceae Exb

ucklandiaceae Exocarpaceae

Fabaceae Fagaceae Falcatifoliaceae Ferulaceae Festucaceae Ficaceae Ficoidaceae Fit

zroyaceae FlacourtiaceaeFlagellariaceae Flindersiaceae Foetidiaceae Forestieraceae

Fothergillaceae Fouquieriaceae Fragariaceae FrancoaceaeFrangulaceae Frankeniacea

e Fraxinaceae Fraxinellaceae Freycinetiaceae Fritillariaceae Fuchsiaceae Fumariacea

eFunkiaceae

Gaiadendraceae Galacaceae Galanthaceae Galaxiaceae Galedupaceae Galeniaceae G

aliaceae Garciniaceae GardeniaceaeGardneriaceae Garryaceae Geissolomataceae Ge

itonoplesiaceae Gelsemiaceae Geniostomaceae GentianaceaeGeonomataceae Geosir

idaceae Geraniaceae Gerrardinaceae Gesneriaceae Gethyllidaceae Gilliesiaceae Gina

lloaceaeGinkgoaceae Gisekiaceae Gladiolaceae Glaucidiaceae Glechomaceae Glinace

ae Globulariaceae Gnaphaliaceae GnetaceaeGoetzeaceae Gomortegaceae Gomphiac

eae Gomphrenaceae Gonystylaceae Goodeniaceae Gordoniaceae GouaniaceaeGoup

iaceae Gramineae Gratiolaceae Grewiaceae Greyiaceae Grielaceae Griseliniaceae Gr

onoviaceae GrossulariaceaeGrubbiaceae Guaiacanaceae Guamatelaceae Guettardac

eae Gunneraceae Gustaviaceae Guttiferae GyrocarpaceaeGyrostemonaceae

Hachetteaceae Haemanthaceae Haemodoraceae Halesiaceae Hallieraceae Halocarpa

ceae Halophilaceae HalophytaceaeHaloragaceae Hamamelidaceae Hanguanaceae H

aptanthaceae Harmandaceae Hebenstretiaceae HectorellaceaeHederaceae Hedyosm

aceae Hedyotidaceae Hedysaraceae Heisteriaceae Heleniaceae Heliamphoraceae H

elianthaceaeHelianthemaceae Helichrysaceae Heliconiaceae Helicteraceae Heliotropi

aceae Helleboraceae Heloniadaceae HelosidaceaeHelwingiaceae Hemerocallidaceae

Hemimeridaceae Hemiphylacaceae Henriqueziaceae Henslowiaceae Hermanniaceae

Hernandiaceae Herniariaceae Herreriaceae Hesperocallidaceae Heterantheraceae He

teropyxidaceae HeterostylaceaeHibbertiaceae Hibiscaceae Hilleriaceae Himantandra

ceae Hippocastanaceae Hippocrateaceae HippomanaceaeHippophaeaceae Hippurida

ceae Hirtellaceae Holacanthaceae Homaliaceae Hopkinsiaceae Hoplestigmataceae H

ordeaceaeHornschuchiaceae Hortensiaceae Hortoniaceae Hostaceae Houstoniaceae

Huaceae Huerteaceae HugoniaceaeHumbertiaceae Humiriaceae Hyacinthaceae Hyd

atellaceae Hydnoraceae Hydrangeaceae Hydrastidaceae HydrillaceaeHydrocharitacea

e Hydrocotylaceae Hydrogetonaceae Hydroleaceae Hydropeltidaceae Hydrophylacac

eae HydrophyllaceaeHydrostachyaceae Hymenocardiaceae Hypecoaceae Hyperanth

eraceae Hypericaceae Hypopityaceae HypoxidaceaeHypseocharitaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta

ceae ImperatoriaceaeInocarpaceae Inulaceae Ionidiaceae Iriarteaceae Iridaceae Irvi

ngiaceae Isnardiaceae Isophysidaceae IteaceaeStrasburgeriaceae Ixiaceae Ixioliriace

ae Ixonanthaceae

Jalapaceae Jamboliferaceae Japonoliriaceae Jasionaceae Jasminiaceae Johnsoniaceae

Joinvilleaceae JuglandaceaeJulianiaceae Juncaceae Juncaginaceae Juniperaceae Jus

siaeaceae Justiciaceae

Kadsuraceae Kaliphoraceae Kalmiaceae Kaniaceae Kiggelariaceae Kingdoniaceae Kin

giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae

Krameriaceae

Labiatae Lacandoniaceae Lachenaliaceae Lacistemataceae Lactoridaceae Lactucaceae

Lagerstroemiaceaee LamiaceaeLampsanaceae Lanariaceae Langsdorffiaceae Lantan

aceae Lapageriaceae Lardizabalaceae Lasiopetalaceae LauraceaeLawsoniaceae Laxm

anniaceae Lecythidaceae Ledaceae Ledocarpaceae Leeaceae Leguminosae Leitneriac

eae LemnaceaeLennoaceae Lentibulariaceae Lentiscaceae Leoniaceae Leonticaceae

Lepidobotryaceae LepidocarpaceaeLepidocaryaceae Lepidocerataceae Leptaulaceae

Leptospermaceae Lepturaceae Lepuropetalaceae LeuctenbergiaceaeLibocedraceae

Licaniaceae Ligustraceae Lilaeaceae Liliaceae Limeaceae Limnanthaceae Limnocharit

aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace

ae Linderniaceae Lindleyaceae Linnaeaceae LippayaceaeLiriaceae Liriodendraceae Li

ssocarpaceae Littorellaceae Loasaceae Lobeliaceaee Loganiaceae LomandraceaeLoni

ceraceae Lopeziaceae Lophiocarpaceae Lophiolaceae Lophiraceae Lophophytaceae L

ophopyxidaceae LoranthaceaeLotaceae Lowiaceae Lupulaceaee Luxemburgiaceae Lu

zuriagaceae Lychnidaceae Lyciaceae LyginiaceaeLygodisodeaceae Lysimachiaceae Ly

thraceae

Macarisiaceae Mackinlayaceae Madiaceae Maesaceae Magnoliaceae Malaceae Male

sherbiaceae MalortieaceaeMalpighiaceae Malvaceae Mangiaceae Manicariaceae Ma

paniaceae Marantaceae Marathraceae MarcgraviaceaeMartyniaceae Mastixiaceae

Matricariaceae Maundiaceae Mayacaceae Medeolaceae Medusagynaceae Peridiscac

eaeMelaleucaceae Melampyraceae Melanophyllaceae Melanthiaceae Melastomatac

eae Meliaceae Melianthaceae MelicaceaeMeliosmaceae Melittidaceae Melochiacea

e Memecylaceae Mendonciaceae Menispermaceae Menthaceae MenyanthaceaeMe

nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

Nypaceae Nyssaceae

Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

ceae Papilionaceae Pappophoraceae PapyraceaeParacryphiaceae Parasitaxaceae Par

ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

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hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

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cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

50

ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 38: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

38

Plocospermataceae [Oleaceae [Tetrachondraceae [Calceolariaceae

[Gesneriaceae Peltanthera [Pedaliaceae Carlemanniaceae Martyniaceae Stilbaceae

Plantaginaceae Linderniaceae Bignoniaceae Verbenaceae Lamiaceae Paulowniaceae

Schlegeliaceae Thomandersiaceae Phrymaceae Rehmannia Acanthaceae

Scrophulariaceae Orobanchaceae Byblidaceae Lentibulariaceae]]]]]

Solanales

[Montiniaceae [Sphenocleaceae + Hydroleaceae]] [Convolvulaceae + Solanaceae]

CAMPANULIDASTERID II

Aquifoliales

[[Cardiopteridaceae + Stemonuraceae] [Aquifoliaceae [Helwingiaceae +

Phyllonomaceae]]

Asterales

[Rousseaceae + Campanulaceae] Pentaphragmataceae [[Alseuosmiaceae [Phellinaceae

+ Argophyllaceae]] Stylidiaceae [Menyanthaceae [Goodeniaceae [Calyceraceae +

Asteraceae]]]]

Escalloniales

Escalloniaceae

Bruniales

[Bruniaceae + Columelliaceae]

Apiales

Pennantiaceae [Torricelliaceae [Griseliniaceae [Pittosporaceae [Araliaceae

[Myodocarpaceae + Apiaceae]]]]]

Paracryphiales

Paracryphiaceae

Dipsacales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

39

Adoxaceae [Diervillaceae [Caprifoliaceae [Linnaeaceae [Morinaceae [Dipsacaceae +

Valerianaceae]]]]]

LINKS TO ORDERS AND FAMILIES

[Back to Top]

MAIN GROUPINGS

Asterids commelinids core eudicots asterid 1 asterid

2 eudicots gymnosperms Magnoliophyta monocots N-fixing claderosids seed plants

ALPHABETICAL LISTING OF ALL ORDINAL NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | L | M | N | O | P | Q | R | S | T | U | V | W | X | Z

Abietales Acanthales Acerales Acorales Actinidiales Actinostrobales Adoxales Aescul

ales Agavales Aizoales AkanialesAlismatales Alliales Alseuosmiales Alstroemeriales

Altingiales Amaranthales Amaryllidales Amborellales AmbrosialesAmmiales Amomal

es Ancistrocladales Anisophylleales Annonales Anthobolales Apiales Apocynales Apo

nogetonalesAquifoliales Arales Araliales Aralidiales Araucariales Arecales Aristolochi

ales Athrotaxidales Asarales AsclepiadalesAsparagales Asphodelales Asparagales Ast

eliales Atriplicales Aucubales Austrobaileyales Avenales

Balanitales Balanopales Balanophorales Balsaminales Barbeyales Barclayales Batales

Begoniales BerberidalesBerberidopsidales Betulales Biebersteiniales Bignoniales Bix

ales Boraginales Brassicales Brexiales Bromeliales BrunialesBrunoniales Burmannial

es Burserales Butomales Buxales Byblidales

Cactales Callitrichales Calycanthales Calycerales Campanulales Campynematales Can

ellales Cannales CapparalesCaprifoliales Cardiopteridales Carduales Caricales Carle

manniales Caryophyllales Cassiales Casuarinales CelastralesCentrolepidales Cephalot

ales Cephalotaxales Ceratophyllales Cercidiphyllales Chenopodiales Chironiales Chlor

anthalesChrysobalanales Cinchonales Circaeasterales Cistales Citrales Cocosales Col

chicales Columelliales CombretalesCommelinales Connarales Convolvulales Coriarial

es Cornales Corylales Corynocarpales Crassulales CrossosomatalesCucurbitales Cunn

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

40

inghamiales Cunoniales Cupressales Cycadales Cyclanthales Cymodoceales Cynarales

CynomorialesCyperales Cyrillales Cytinales

Daphnales Daphniphyllales Dasypogonales Datiscales Degeneriales Desfontainiales D

iapensiales Didymelales DillenialesDioales Dioncophyllales Dioscoreales Diospyrales

Dipentodontales Dipsacales Droserales

Ebenales Echiales Elaeagnales Elaeocarpales Elatinales Elodeales Emmotales Empetr

ales Ephedrales Ericales EriocaulalesErythropalaless Escalloniales Eucommiales Euph

orbiales Eupomatiales Eupteleales Euryalaless

Fabales Fagales Falcatifoliales Ficales Flacourtiales Flagellariales Fouquieriales Franc

oales Frangulales

Galiales Garryales Geissolomatales Gentianales Geraniales Gesneriales Ginkgoales G

laucidiales Globulariales GnetalesGoodeniales Greyiales Griseliniales Grossulariales

Grubbiales Gunnerales Gyrocarpales Gyrostemonales

Haemodorales Haloragales Hamamelidales Hanguanales Heisteriales Helleborales He

lwingiales Himantandrales HippuridalesHomaliales Hortensiales Huales Huerteales

Hydatellales Hydnorales Hydrangeales Hydrastidales HydrocharitalesHydropeltidales

Hydrostachyales Hypericales Hypoxidales

Icacinales Illiciales Iridales Irvingales Iteales Ixiales

Jasminales Juglandales Julianiales Juncaginales Juncales

Lacistematales Lactoridales Lamiales Lardizabalales Laurales Lecythidales Ledocarpal

es Leitneriales LentibularialesLigustrales Liliales Limnanthales Linales Liriales Loasal

es Lobeliales Loganiales Lonicerales Loranthales LowialesLythrales

Magnoliales Malpighiales Malvales Marathrales Marcgraviales Mayacales Medusagy

nales Medusandrales MelanthialesMelastomatales Meliales Melianthales Meliosmal

es Menispermales Menyanthales Metteniusales MitrastemonalesMiyoshiales Moni

miales Moringales Musales Myricales Myristicales Myrothamnales Myrsinales Myrta

les

Najadales Nandinales Narcissales Nartheciales Nelumbonales Nepenthales Neuradal

es Nitrariales Nolanales NothofagalesNyctaginales Nymphaeales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

41

Ochnales Oenotherales Olacales Oleales Onagrales Oncothecales Opuntiales Orchid

ales Oxalidales

Saxifragales Pandales Pandanales Papaverales Paracryphiales Parasitaxales Paridales

Parnassiales PassifloralesPenaeales Pennantiales Pentaphragmatales Peridiscales P

etiveriales Petrosaviales Phellinales Philydrales PhyllanthalesPhysenales Phytolaccal

es Picramniales Pinales Pinguiculales Piperales Pittosporales Plantaginales Platanale

sPlumbaginales Poales Podocarpales Podophyllales Podostemales Polemoniales Pol

ygalales Polygonales PontederialesPortulacales Posidoniales Potamogetonales Primu

lales Proteales

Quercales Quillajales Quintiniales

Rafflesiales Ranunculales Rapateales Resedales Restionales Rhabdodendrales Rhamn

ales Rhinanthales RhizophoralesRhodorales Rhoipteleales Roridulales Rosales Rouss

eales Rubiales Ruppiales Rutales

Sabiales Salicales Salvadorales Samolales Samydales Sanguisorbales Santalales Sapin

dales Sapotales SarracenialesSaxegotheales Saxifragales Scheuchzeriales Sciadopitya

les Scleranthales Scrophulariales Scyphostegiales SedalesSilenales Simmondsiales S

milacales Solanales Sphenocleales Sphenostemonales Stangeriales Stellariales Stemo

nalesStilbales Stylidiales Styracales Surianales

Taccales Tamales Tamaricales Taxales Taxodiales Tecophilaeales Terebinthales Tern

stroemiales Theales TheligonalesThymelaeales Tiliales Tofieldiales Torricelliales Tov

ariales Tribelales Trilliales Trimeniales Triuridales TrochodendralesTropaeolales Turn

erales Typhales

Ulmales Urticales

Vacciniales Vahliales Vallisneriales Velloziales Veratrales Verbenales Viburnales Vinc

ales Violales Viscales VitalesVochysiales

Welwitschiales Winterales

Xanthorrhoeales Ximeniales Xyridales

Zamiales Zingiberales Zosterales Zygophyllales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

42

ALPHABETICAL LISTING OF ALL FAMILY NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | K | L | M | N | O | P | Q | R | S | T | U | V | W | X | Y

|Z

Back to Home

In some cases clicking on a family synonym will send you directly to the subfamily in

which the genus of the basionym is to be found

Abietaceae Abolbodaceae Abrophyllaceae Acalyphaceae Acanthaceae [Acanthochla

mydaceae] Acarnaceaee AceraceaeAchariaceae Achatocarpaceae Achradaceae Achy

ranthaceae Acmopylaceae Acoraceae Acristaceae ActaeaceaeActinidiaceae Actinota

ceae Amentotaxaceae Adenogrammaceae Adoxaceae Aegialitidaceae Aegicerataceae

Aegilopaceae Aeginetiaceaee Aegiphilaceae Aesculaceae Aextoxicaceae Agapanthac

eae Agavaceae AgdestidaceaeAgialidaceae Agrimoniaceae Agrostidaceae Ailanthace

ae Aitoniaceae Aizoaceae Akaniaceae Alangiaceae AlchemillaceaeAldrovandaceae A

lismataceae Alliaceae Allioniaceae Allophylaceae Aloaceae Alopecuraceae Alpiniacea

e AlseuosmiaceaeAlsinaceae Alsodeiaceae Alstroemeriaceae Altingiaceae Alzateacea

e Amaranthaceae Amaryllidaceae Amborellaceae Ambrosiaceae Amentotaxaceae A

mmanniaceae Ammiaceae Amomaceae Ampelidopsaceae Amygdalaceae Amyridacea

eAnacardiaceae Anagallidaceae Anarthriaceae Anchusaceae Ancistrocladaceae Andr

omedaceae AndropogonaceaeAndrostachyaceae Androsynaceae Anemarrhenaceae

Anemonaceae Angelicaceae Anisophylleaceae AnnonaceaeAnomochloaceae Anopter

aceae Anrederaceae Anthemidaceae Anthericaceae Anthobolaceae Antidesmataceae

Antirrhinaceae Antoniaceae Aparinaceae Aphanopetalaceae Aphloiaceae Aphyllanth

aceae Apiaceae ApocynaceaeApodanthaceae Aponogetonaceae Aporusaceae Apose

ridaceae Apostasiaceae Aptandraceae AquifoliaceaeAquilariaceae Aquilegiaceae A

raceae Aragoaceae Araliaceae Aralidiaceaee Araucariaceae Arbutaceae Arceuthidace

aeArceuthobiaceae Arctostaphylaceae Arctotidaceae Ardisiaceae Arecaceae Argoph

yllaceae Arisaraceae AristolochiaceaeAristoteliaceae Arjonaceae Armeriaceae Artem

isiaceae Arthrotaxidaceae Artocarpaceae Arundinaceae ArundinellaceaeAsaraceae A

sclepiadaceae Ascyraceaee Asiraceae Asparagaceae Asperulaceae Asphodelaceae As

pidistraceaeeAsteliaceae Asteraceae Asteranthaceae Asteropeiaceae Astragalaceae

Astrocarpaceae AthanasiaceaeAtherospermataceae Atriplicaceae Atropaceae Aucuba

ceae Aurantiaceae Austrobaileyaceae Austrotaxaceae AvenaceaeAverrhoaceae Avet

raceae Avicenniaceae Azaleaceae Azimaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

43

Balanitaceae Balanopaceae Balanophoraceae Balsameaceae Balsaminaceae Bambusa

ceae Barbaceniaceae BarbeuiaceaeBarbeyaceae Barclayaceae Barringtoniaceae Base

llaceae Bataceae Baueraceae Bauhiniaceae Baxteriaceae BegoniaceaeBehniaceae B

elangeraceae Belloniaceae Bembiciaceae Berberidaceae Berberidopsidaceae Berryac

eaee BersamaceaeBertyaceae Berzeliaceae Besleriaceae Betaceae Betulaceae Biebe

rsteiniaceae Bifariaceae Bignoniaceae BischofiaceaeBixaceae Blakeaceae Blandfordia

ceae Blattiaceae Blepharocaryaceae Blitaceae Blyxaceae Boerlagellaceae Bolivaracea

eBombacaceae Bonnetiaceae Bontiaceae Boopidaceae Boraginaceae Borassaceae B

oroniaceae BoryaceaeBotryodendraceae Bougainvilleaceae Boweniaceae Brassicacea

e Brachycaulaceae Bretschneideraceae BrexiaceaeBromeliaceae Brownlowiaceae Br

unelliaceae Bruniaceae Brunoniaceae Brunsvigiaceae Bryoniaceae BuchneraceaeBuci

daceae Buddlejaceae Buglossaceae Bulbocodiaceae Bumeliaceae Bupleuraceae Burc

hardiaceae BurmanniaceaeBurseraceae Butneriaceae Butomaceae Buxaceae Byblida

ceae Byttneriaceae

Cabombaceae Cacaoaceae Cactaceae Caesalpiniaceae Caladiaceae Calamaceae Calce

olariaceae CalectasiaceaeCalendulaceae Callaceae Callicomaceae Calligonaceae Calli

traceae Callitrichaceae Calochortaceae CalophyllaceaeCalthaceae Calycanthaceae Ca

lyceraceae Cambogiaceae Camelliaceae Campanulaceae CampynemataceaeCanacom

yricaceae Canellaceae Cannabaceae Cannaceae Canopodaceae Canotiaceae Cansjera

ceae CapparaceaeCaprariaceae Caprifoliaceae Cardiopteridaceae Carduaceae Carica

ceae Carissaceae Carlemanniaceae CarpinaceaeCarpodetaceae Cartonemataceae Ca

ryocaraceae Caryophyllaceae Casearieaceae Cassiaceae CassipoureaceaeCassythacea

e Castaneaceae Castelaceae Casuarinaceae Catesbaeaceae Cathedraceae Cecropiace

ae CedraceaeCedrelaceae Celastraceae Celosiaceae Celtidaceae Centaureaceae Cen

trolepidaceae Centroplacaceae CepaceaeCephalanthaceae Cephalotaceae Cephalota

xaceae Cerastiaceae Ceratoniaceae Ceratophyllaceae CerberaceaeCercidiphyllaceae

Cercocarpaceae Cercodiaceae Cereaceae Cerinthaceae Ceroxylaceae Cestraceae Cev

alliaceaeChailletiaceae Chamaedoraceae Chamaemoraceae Chamelauciaceae Chaun

ochitonaceae Chelidoniaceae ChelonaceaeChenopodiaceae Chimonanthaceae Chingi

thamnaceae Chionographidaceae Chiranthodendraceae ChironiaceaeChlaenaceae Ch

loanthaceae Chloranthaceae Chloridaceae Chlorogalaceae Chrysobalanaceae Cicerac

eae CichoriaceaeCimicifugaceae Cinchoniaceae Circaeaceae Circaeasteraceae Cissac

eae Cistaceae Citraceae Clematidaceae CleomaceaeClethraceae Clusiaceae Cneorac

eae Cnestidaceae Cnicaceae Cobaeaceae Cochlospermaceae Cocosaceae Coffeaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

44

Colchicaceae Coleogynaceae Columelliaceae Combretaceae Commelinaceae Comocl

adiaceae Compositae CompsoaceaeConantheraceae Connaraceae Conostylidaceae C

onvallariaceae Convolvulaceae Cordiaceae CoreopsidaceaeCoriandraceae Coriariacea

e Coridaceae Corispermaceae Cornaceae Corokiaceae Coronillaceae Correaceae Cor

rigiolaceaeCorsiaceae Corydalaceae Corylaceae Corynocarpaceae Coryphaceae Cost

aceae Cotyledonaceae CoulaceaeCoutariaceae Coutoubiaceae Crassulaceae Crescen

tiaceae Cressaceae Crinaceae Crocaceae CroomiaceaeCrossosomataceae Crotonacea

e Cruciferae Cryptaceae Crypteroniaceae Cryptocorynaceae CryptomeriaceaeCtenol

ophonaceae Cucurbitaceae Cunninghamiaceae Cunoniaceae Cupressaceae Curcumac

eae Curtisiaceae CuscutaceaeCyananthaceae Cyanastraceae Cynanchaceae Cyanella

ceae Cycadaceae Cyclanthaceae CyclantheraceaeCyclocheilaceae Cydoniaceae Cymo

doceaceae Cynaraceae Cynocrambaceae Cynomoriaceae Cyperaceae CyphiaceaeCyp

hocarpaceae Cypripediaceae Cyrillaceae Cyrtandraceae Cyrtanthaceae Cytinaceae

Dacrycarpaceae Dactylanthaceae Damasoniaceae Daphnaceae Daphniphyllaceae Das

ypogonaceae DatiscaceaeDaturaceae Daucaceae Davidiaceae Davidsoniaceae Decais

neaceae Deeringiaceae Degeneriaceae DelphiniaceaeDendrophthoaceae Columelliac

eae Detariaceae Dialypetalanthaceae Dianellaceae Dianthaceae DiapensiaceaeDicha

petalaceae Dichondraceae Diclidantheraceae Dicrastylidiaceae Dictamnaceae Didiere

aceae DidymelaceaeDidymocarpaceae Diegodendraceae Diervillaceae Digitalidaceae

e Dilatridaceae Dilleniaceae Dioaceae DionaeaceaeDioncophyllaceae Dioscoreaceae

Diosmaceae Diospyraceae Dipentodontaceae Diphylleiaceae Diplolaenaceae Dipsaca

ceaeDipterocarpaceae Dirachmaceae Disanthaceae Diselmaceae Dodonaeaceae Do

mbeyaceae Donatiaceae DorsteniaceaeDortmannaceae Doryanthaceae Drabaceae D

racaenaceae Dracontiaceae Drimyidaceae Droseraceae DrosophyllaceaeDryadaceae

Duabangaceae Duckeodendraceae Dulongiaceae Durantaceae Durionaceae Dysphani

aceae

Ebenaceae Eccremidaceae Ecdeiocoleaceae Echiaceae Echinopaceae Ehretiaceae Ela

eagnaceae ElaeocarpaceaeElegiaceae Elatinaceae Ellisiaceae Ellisiophyllaceae Elodea

ceae Elytranthaceae Embeliaceae Emblingiaceae EmmotaceaeEmpetraceae Encepha

lartaceae Engelhardtiaceae Enhalaceae Epacridaceae Ephedraceae Ephemeranthacea

eEpilobiaceae Epimediaceae Eragrostidaceae Eremolepidaceae Eremosynaceae Erica

ceae Erinaceae EriocaulaceaeEriogonaceae Eriospermaceae Erodiaceae Erycibaceae

Eryngiaceae Erysimaceae Erythroniaceae ErythropalaceaeErythrospermaceae Erythro

xylaceae Escalloniaceae Eschscholziaceae Eucomidaceae Eucommiaceae Eucryphiace

aeEuonymaceae Eupatoriaceae Euphorbiaceae Euphrasiaceae Euphroniaceae Eupo

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

45

matiaceae Eupteleaceae EuryalaceaeEustrephaceae Euthemidaceae Eutocaceae Exb

ucklandiaceae Exocarpaceae

Fabaceae Fagaceae Falcatifoliaceae Ferulaceae Festucaceae Ficaceae Ficoidaceae Fit

zroyaceae FlacourtiaceaeFlagellariaceae Flindersiaceae Foetidiaceae Forestieraceae

Fothergillaceae Fouquieriaceae Fragariaceae FrancoaceaeFrangulaceae Frankeniacea

e Fraxinaceae Fraxinellaceae Freycinetiaceae Fritillariaceae Fuchsiaceae Fumariacea

eFunkiaceae

Gaiadendraceae Galacaceae Galanthaceae Galaxiaceae Galedupaceae Galeniaceae G

aliaceae Garciniaceae GardeniaceaeGardneriaceae Garryaceae Geissolomataceae Ge

itonoplesiaceae Gelsemiaceae Geniostomaceae GentianaceaeGeonomataceae Geosir

idaceae Geraniaceae Gerrardinaceae Gesneriaceae Gethyllidaceae Gilliesiaceae Gina

lloaceaeGinkgoaceae Gisekiaceae Gladiolaceae Glaucidiaceae Glechomaceae Glinace

ae Globulariaceae Gnaphaliaceae GnetaceaeGoetzeaceae Gomortegaceae Gomphiac

eae Gomphrenaceae Gonystylaceae Goodeniaceae Gordoniaceae GouaniaceaeGoup

iaceae Gramineae Gratiolaceae Grewiaceae Greyiaceae Grielaceae Griseliniaceae Gr

onoviaceae GrossulariaceaeGrubbiaceae Guaiacanaceae Guamatelaceae Guettardac

eae Gunneraceae Gustaviaceae Guttiferae GyrocarpaceaeGyrostemonaceae

Hachetteaceae Haemanthaceae Haemodoraceae Halesiaceae Hallieraceae Halocarpa

ceae Halophilaceae HalophytaceaeHaloragaceae Hamamelidaceae Hanguanaceae H

aptanthaceae Harmandaceae Hebenstretiaceae HectorellaceaeHederaceae Hedyosm

aceae Hedyotidaceae Hedysaraceae Heisteriaceae Heleniaceae Heliamphoraceae H

elianthaceaeHelianthemaceae Helichrysaceae Heliconiaceae Helicteraceae Heliotropi

aceae Helleboraceae Heloniadaceae HelosidaceaeHelwingiaceae Hemerocallidaceae

Hemimeridaceae Hemiphylacaceae Henriqueziaceae Henslowiaceae Hermanniaceae

Hernandiaceae Herniariaceae Herreriaceae Hesperocallidaceae Heterantheraceae He

teropyxidaceae HeterostylaceaeHibbertiaceae Hibiscaceae Hilleriaceae Himantandra

ceae Hippocastanaceae Hippocrateaceae HippomanaceaeHippophaeaceae Hippurida

ceae Hirtellaceae Holacanthaceae Homaliaceae Hopkinsiaceae Hoplestigmataceae H

ordeaceaeHornschuchiaceae Hortensiaceae Hortoniaceae Hostaceae Houstoniaceae

Huaceae Huerteaceae HugoniaceaeHumbertiaceae Humiriaceae Hyacinthaceae Hyd

atellaceae Hydnoraceae Hydrangeaceae Hydrastidaceae HydrillaceaeHydrocharitacea

e Hydrocotylaceae Hydrogetonaceae Hydroleaceae Hydropeltidaceae Hydrophylacac

eae HydrophyllaceaeHydrostachyaceae Hymenocardiaceae Hypecoaceae Hyperanth

eraceae Hypericaceae Hypopityaceae HypoxidaceaeHypseocharitaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

46

Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta

ceae ImperatoriaceaeInocarpaceae Inulaceae Ionidiaceae Iriarteaceae Iridaceae Irvi

ngiaceae Isnardiaceae Isophysidaceae IteaceaeStrasburgeriaceae Ixiaceae Ixioliriace

ae Ixonanthaceae

Jalapaceae Jamboliferaceae Japonoliriaceae Jasionaceae Jasminiaceae Johnsoniaceae

Joinvilleaceae JuglandaceaeJulianiaceae Juncaceae Juncaginaceae Juniperaceae Jus

siaeaceae Justiciaceae

Kadsuraceae Kaliphoraceae Kalmiaceae Kaniaceae Kiggelariaceae Kingdoniaceae Kin

giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae

Krameriaceae

Labiatae Lacandoniaceae Lachenaliaceae Lacistemataceae Lactoridaceae Lactucaceae

Lagerstroemiaceaee LamiaceaeLampsanaceae Lanariaceae Langsdorffiaceae Lantan

aceae Lapageriaceae Lardizabalaceae Lasiopetalaceae LauraceaeLawsoniaceae Laxm

anniaceae Lecythidaceae Ledaceae Ledocarpaceae Leeaceae Leguminosae Leitneriac

eae LemnaceaeLennoaceae Lentibulariaceae Lentiscaceae Leoniaceae Leonticaceae

Lepidobotryaceae LepidocarpaceaeLepidocaryaceae Lepidocerataceae Leptaulaceae

Leptospermaceae Lepturaceae Lepuropetalaceae LeuctenbergiaceaeLibocedraceae

Licaniaceae Ligustraceae Lilaeaceae Liliaceae Limeaceae Limnanthaceae Limnocharit

aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace

ae Linderniaceae Lindleyaceae Linnaeaceae LippayaceaeLiriaceae Liriodendraceae Li

ssocarpaceae Littorellaceae Loasaceae Lobeliaceaee Loganiaceae LomandraceaeLoni

ceraceae Lopeziaceae Lophiocarpaceae Lophiolaceae Lophiraceae Lophophytaceae L

ophopyxidaceae LoranthaceaeLotaceae Lowiaceae Lupulaceaee Luxemburgiaceae Lu

zuriagaceae Lychnidaceae Lyciaceae LyginiaceaeLygodisodeaceae Lysimachiaceae Ly

thraceae

Macarisiaceae Mackinlayaceae Madiaceae Maesaceae Magnoliaceae Malaceae Male

sherbiaceae MalortieaceaeMalpighiaceae Malvaceae Mangiaceae Manicariaceae Ma

paniaceae Marantaceae Marathraceae MarcgraviaceaeMartyniaceae Mastixiaceae

Matricariaceae Maundiaceae Mayacaceae Medeolaceae Medusagynaceae Peridiscac

eaeMelaleucaceae Melampyraceae Melanophyllaceae Melanthiaceae Melastomatac

eae Meliaceae Melianthaceae MelicaceaeMeliosmaceae Melittidaceae Melochiacea

e Memecylaceae Mendonciaceae Menispermaceae Menthaceae MenyanthaceaeMe

nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

47

Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

Nypaceae Nyssaceae

Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

ceae Papilionaceae Pappophoraceae PapyraceaeParacryphiaceae Parasitaxaceae Par

ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

48

hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

49

cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

50

ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 39: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

39

Adoxaceae [Diervillaceae [Caprifoliaceae [Linnaeaceae [Morinaceae [Dipsacaceae +

Valerianaceae]]]]]

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MAIN GROUPINGS

Asterids commelinids core eudicots asterid 1 asterid

2 eudicots gymnosperms Magnoliophyta monocots N-fixing claderosids seed plants

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Abietales Acanthales Acerales Acorales Actinidiales Actinostrobales Adoxales Aescul

ales Agavales Aizoales AkanialesAlismatales Alliales Alseuosmiales Alstroemeriales

Altingiales Amaranthales Amaryllidales Amborellales AmbrosialesAmmiales Amomal

es Ancistrocladales Anisophylleales Annonales Anthobolales Apiales Apocynales Apo

nogetonalesAquifoliales Arales Araliales Aralidiales Araucariales Arecales Aristolochi

ales Athrotaxidales Asarales AsclepiadalesAsparagales Asphodelales Asparagales Ast

eliales Atriplicales Aucubales Austrobaileyales Avenales

Balanitales Balanopales Balanophorales Balsaminales Barbeyales Barclayales Batales

Begoniales BerberidalesBerberidopsidales Betulales Biebersteiniales Bignoniales Bix

ales Boraginales Brassicales Brexiales Bromeliales BrunialesBrunoniales Burmannial

es Burserales Butomales Buxales Byblidales

Cactales Callitrichales Calycanthales Calycerales Campanulales Campynematales Can

ellales Cannales CapparalesCaprifoliales Cardiopteridales Carduales Caricales Carle

manniales Caryophyllales Cassiales Casuarinales CelastralesCentrolepidales Cephalot

ales Cephalotaxales Ceratophyllales Cercidiphyllales Chenopodiales Chironiales Chlor

anthalesChrysobalanales Cinchonales Circaeasterales Cistales Citrales Cocosales Col

chicales Columelliales CombretalesCommelinales Connarales Convolvulales Coriarial

es Cornales Corylales Corynocarpales Crassulales CrossosomatalesCucurbitales Cunn

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

40

inghamiales Cunoniales Cupressales Cycadales Cyclanthales Cymodoceales Cynarales

CynomorialesCyperales Cyrillales Cytinales

Daphnales Daphniphyllales Dasypogonales Datiscales Degeneriales Desfontainiales D

iapensiales Didymelales DillenialesDioales Dioncophyllales Dioscoreales Diospyrales

Dipentodontales Dipsacales Droserales

Ebenales Echiales Elaeagnales Elaeocarpales Elatinales Elodeales Emmotales Empetr

ales Ephedrales Ericales EriocaulalesErythropalaless Escalloniales Eucommiales Euph

orbiales Eupomatiales Eupteleales Euryalaless

Fabales Fagales Falcatifoliales Ficales Flacourtiales Flagellariales Fouquieriales Franc

oales Frangulales

Galiales Garryales Geissolomatales Gentianales Geraniales Gesneriales Ginkgoales G

laucidiales Globulariales GnetalesGoodeniales Greyiales Griseliniales Grossulariales

Grubbiales Gunnerales Gyrocarpales Gyrostemonales

Haemodorales Haloragales Hamamelidales Hanguanales Heisteriales Helleborales He

lwingiales Himantandrales HippuridalesHomaliales Hortensiales Huales Huerteales

Hydatellales Hydnorales Hydrangeales Hydrastidales HydrocharitalesHydropeltidales

Hydrostachyales Hypericales Hypoxidales

Icacinales Illiciales Iridales Irvingales Iteales Ixiales

Jasminales Juglandales Julianiales Juncaginales Juncales

Lacistematales Lactoridales Lamiales Lardizabalales Laurales Lecythidales Ledocarpal

es Leitneriales LentibularialesLigustrales Liliales Limnanthales Linales Liriales Loasal

es Lobeliales Loganiales Lonicerales Loranthales LowialesLythrales

Magnoliales Malpighiales Malvales Marathrales Marcgraviales Mayacales Medusagy

nales Medusandrales MelanthialesMelastomatales Meliales Melianthales Meliosmal

es Menispermales Menyanthales Metteniusales MitrastemonalesMiyoshiales Moni

miales Moringales Musales Myricales Myristicales Myrothamnales Myrsinales Myrta

les

Najadales Nandinales Narcissales Nartheciales Nelumbonales Nepenthales Neuradal

es Nitrariales Nolanales NothofagalesNyctaginales Nymphaeales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

41

Ochnales Oenotherales Olacales Oleales Onagrales Oncothecales Opuntiales Orchid

ales Oxalidales

Saxifragales Pandales Pandanales Papaverales Paracryphiales Parasitaxales Paridales

Parnassiales PassifloralesPenaeales Pennantiales Pentaphragmatales Peridiscales P

etiveriales Petrosaviales Phellinales Philydrales PhyllanthalesPhysenales Phytolaccal

es Picramniales Pinales Pinguiculales Piperales Pittosporales Plantaginales Platanale

sPlumbaginales Poales Podocarpales Podophyllales Podostemales Polemoniales Pol

ygalales Polygonales PontederialesPortulacales Posidoniales Potamogetonales Primu

lales Proteales

Quercales Quillajales Quintiniales

Rafflesiales Ranunculales Rapateales Resedales Restionales Rhabdodendrales Rhamn

ales Rhinanthales RhizophoralesRhodorales Rhoipteleales Roridulales Rosales Rouss

eales Rubiales Ruppiales Rutales

Sabiales Salicales Salvadorales Samolales Samydales Sanguisorbales Santalales Sapin

dales Sapotales SarracenialesSaxegotheales Saxifragales Scheuchzeriales Sciadopitya

les Scleranthales Scrophulariales Scyphostegiales SedalesSilenales Simmondsiales S

milacales Solanales Sphenocleales Sphenostemonales Stangeriales Stellariales Stemo

nalesStilbales Stylidiales Styracales Surianales

Taccales Tamales Tamaricales Taxales Taxodiales Tecophilaeales Terebinthales Tern

stroemiales Theales TheligonalesThymelaeales Tiliales Tofieldiales Torricelliales Tov

ariales Tribelales Trilliales Trimeniales Triuridales TrochodendralesTropaeolales Turn

erales Typhales

Ulmales Urticales

Vacciniales Vahliales Vallisneriales Velloziales Veratrales Verbenales Viburnales Vinc

ales Violales Viscales VitalesVochysiales

Welwitschiales Winterales

Xanthorrhoeales Ximeniales Xyridales

Zamiales Zingiberales Zosterales Zygophyllales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

42

ALPHABETICAL LISTING OF ALL FAMILY NAMES OF SEED PLANTS WITH LINKS

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|Z

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In some cases clicking on a family synonym will send you directly to the subfamily in

which the genus of the basionym is to be found

Abietaceae Abolbodaceae Abrophyllaceae Acalyphaceae Acanthaceae [Acanthochla

mydaceae] Acarnaceaee AceraceaeAchariaceae Achatocarpaceae Achradaceae Achy

ranthaceae Acmopylaceae Acoraceae Acristaceae ActaeaceaeActinidiaceae Actinota

ceae Amentotaxaceae Adenogrammaceae Adoxaceae Aegialitidaceae Aegicerataceae

Aegilopaceae Aeginetiaceaee Aegiphilaceae Aesculaceae Aextoxicaceae Agapanthac

eae Agavaceae AgdestidaceaeAgialidaceae Agrimoniaceae Agrostidaceae Ailanthace

ae Aitoniaceae Aizoaceae Akaniaceae Alangiaceae AlchemillaceaeAldrovandaceae A

lismataceae Alliaceae Allioniaceae Allophylaceae Aloaceae Alopecuraceae Alpiniacea

e AlseuosmiaceaeAlsinaceae Alsodeiaceae Alstroemeriaceae Altingiaceae Alzateacea

e Amaranthaceae Amaryllidaceae Amborellaceae Ambrosiaceae Amentotaxaceae A

mmanniaceae Ammiaceae Amomaceae Ampelidopsaceae Amygdalaceae Amyridacea

eAnacardiaceae Anagallidaceae Anarthriaceae Anchusaceae Ancistrocladaceae Andr

omedaceae AndropogonaceaeAndrostachyaceae Androsynaceae Anemarrhenaceae

Anemonaceae Angelicaceae Anisophylleaceae AnnonaceaeAnomochloaceae Anopter

aceae Anrederaceae Anthemidaceae Anthericaceae Anthobolaceae Antidesmataceae

Antirrhinaceae Antoniaceae Aparinaceae Aphanopetalaceae Aphloiaceae Aphyllanth

aceae Apiaceae ApocynaceaeApodanthaceae Aponogetonaceae Aporusaceae Apose

ridaceae Apostasiaceae Aptandraceae AquifoliaceaeAquilariaceae Aquilegiaceae A

raceae Aragoaceae Araliaceae Aralidiaceaee Araucariaceae Arbutaceae Arceuthidace

aeArceuthobiaceae Arctostaphylaceae Arctotidaceae Ardisiaceae Arecaceae Argoph

yllaceae Arisaraceae AristolochiaceaeAristoteliaceae Arjonaceae Armeriaceae Artem

isiaceae Arthrotaxidaceae Artocarpaceae Arundinaceae ArundinellaceaeAsaraceae A

sclepiadaceae Ascyraceaee Asiraceae Asparagaceae Asperulaceae Asphodelaceae As

pidistraceaeeAsteliaceae Asteraceae Asteranthaceae Asteropeiaceae Astragalaceae

Astrocarpaceae AthanasiaceaeAtherospermataceae Atriplicaceae Atropaceae Aucuba

ceae Aurantiaceae Austrobaileyaceae Austrotaxaceae AvenaceaeAverrhoaceae Avet

raceae Avicenniaceae Azaleaceae Azimaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Balanitaceae Balanopaceae Balanophoraceae Balsameaceae Balsaminaceae Bambusa

ceae Barbaceniaceae BarbeuiaceaeBarbeyaceae Barclayaceae Barringtoniaceae Base

llaceae Bataceae Baueraceae Bauhiniaceae Baxteriaceae BegoniaceaeBehniaceae B

elangeraceae Belloniaceae Bembiciaceae Berberidaceae Berberidopsidaceae Berryac

eaee BersamaceaeBertyaceae Berzeliaceae Besleriaceae Betaceae Betulaceae Biebe

rsteiniaceae Bifariaceae Bignoniaceae BischofiaceaeBixaceae Blakeaceae Blandfordia

ceae Blattiaceae Blepharocaryaceae Blitaceae Blyxaceae Boerlagellaceae Bolivaracea

eBombacaceae Bonnetiaceae Bontiaceae Boopidaceae Boraginaceae Borassaceae B

oroniaceae BoryaceaeBotryodendraceae Bougainvilleaceae Boweniaceae Brassicacea

e Brachycaulaceae Bretschneideraceae BrexiaceaeBromeliaceae Brownlowiaceae Br

unelliaceae Bruniaceae Brunoniaceae Brunsvigiaceae Bryoniaceae BuchneraceaeBuci

daceae Buddlejaceae Buglossaceae Bulbocodiaceae Bumeliaceae Bupleuraceae Burc

hardiaceae BurmanniaceaeBurseraceae Butneriaceae Butomaceae Buxaceae Byblida

ceae Byttneriaceae

Cabombaceae Cacaoaceae Cactaceae Caesalpiniaceae Caladiaceae Calamaceae Calce

olariaceae CalectasiaceaeCalendulaceae Callaceae Callicomaceae Calligonaceae Calli

traceae Callitrichaceae Calochortaceae CalophyllaceaeCalthaceae Calycanthaceae Ca

lyceraceae Cambogiaceae Camelliaceae Campanulaceae CampynemataceaeCanacom

yricaceae Canellaceae Cannabaceae Cannaceae Canopodaceae Canotiaceae Cansjera

ceae CapparaceaeCaprariaceae Caprifoliaceae Cardiopteridaceae Carduaceae Carica

ceae Carissaceae Carlemanniaceae CarpinaceaeCarpodetaceae Cartonemataceae Ca

ryocaraceae Caryophyllaceae Casearieaceae Cassiaceae CassipoureaceaeCassythacea

e Castaneaceae Castelaceae Casuarinaceae Catesbaeaceae Cathedraceae Cecropiace

ae CedraceaeCedrelaceae Celastraceae Celosiaceae Celtidaceae Centaureaceae Cen

trolepidaceae Centroplacaceae CepaceaeCephalanthaceae Cephalotaceae Cephalota

xaceae Cerastiaceae Ceratoniaceae Ceratophyllaceae CerberaceaeCercidiphyllaceae

Cercocarpaceae Cercodiaceae Cereaceae Cerinthaceae Ceroxylaceae Cestraceae Cev

alliaceaeChailletiaceae Chamaedoraceae Chamaemoraceae Chamelauciaceae Chaun

ochitonaceae Chelidoniaceae ChelonaceaeChenopodiaceae Chimonanthaceae Chingi

thamnaceae Chionographidaceae Chiranthodendraceae ChironiaceaeChlaenaceae Ch

loanthaceae Chloranthaceae Chloridaceae Chlorogalaceae Chrysobalanaceae Cicerac

eae CichoriaceaeCimicifugaceae Cinchoniaceae Circaeaceae Circaeasteraceae Cissac

eae Cistaceae Citraceae Clematidaceae CleomaceaeClethraceae Clusiaceae Cneorac

eae Cnestidaceae Cnicaceae Cobaeaceae Cochlospermaceae Cocosaceae Coffeaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Colchicaceae Coleogynaceae Columelliaceae Combretaceae Commelinaceae Comocl

adiaceae Compositae CompsoaceaeConantheraceae Connaraceae Conostylidaceae C

onvallariaceae Convolvulaceae Cordiaceae CoreopsidaceaeCoriandraceae Coriariacea

e Coridaceae Corispermaceae Cornaceae Corokiaceae Coronillaceae Correaceae Cor

rigiolaceaeCorsiaceae Corydalaceae Corylaceae Corynocarpaceae Coryphaceae Cost

aceae Cotyledonaceae CoulaceaeCoutariaceae Coutoubiaceae Crassulaceae Crescen

tiaceae Cressaceae Crinaceae Crocaceae CroomiaceaeCrossosomataceae Crotonacea

e Cruciferae Cryptaceae Crypteroniaceae Cryptocorynaceae CryptomeriaceaeCtenol

ophonaceae Cucurbitaceae Cunninghamiaceae Cunoniaceae Cupressaceae Curcumac

eae Curtisiaceae CuscutaceaeCyananthaceae Cyanastraceae Cynanchaceae Cyanella

ceae Cycadaceae Cyclanthaceae CyclantheraceaeCyclocheilaceae Cydoniaceae Cymo

doceaceae Cynaraceae Cynocrambaceae Cynomoriaceae Cyperaceae CyphiaceaeCyp

hocarpaceae Cypripediaceae Cyrillaceae Cyrtandraceae Cyrtanthaceae Cytinaceae

Dacrycarpaceae Dactylanthaceae Damasoniaceae Daphnaceae Daphniphyllaceae Das

ypogonaceae DatiscaceaeDaturaceae Daucaceae Davidiaceae Davidsoniaceae Decais

neaceae Deeringiaceae Degeneriaceae DelphiniaceaeDendrophthoaceae Columelliac

eae Detariaceae Dialypetalanthaceae Dianellaceae Dianthaceae DiapensiaceaeDicha

petalaceae Dichondraceae Diclidantheraceae Dicrastylidiaceae Dictamnaceae Didiere

aceae DidymelaceaeDidymocarpaceae Diegodendraceae Diervillaceae Digitalidaceae

e Dilatridaceae Dilleniaceae Dioaceae DionaeaceaeDioncophyllaceae Dioscoreaceae

Diosmaceae Diospyraceae Dipentodontaceae Diphylleiaceae Diplolaenaceae Dipsaca

ceaeDipterocarpaceae Dirachmaceae Disanthaceae Diselmaceae Dodonaeaceae Do

mbeyaceae Donatiaceae DorsteniaceaeDortmannaceae Doryanthaceae Drabaceae D

racaenaceae Dracontiaceae Drimyidaceae Droseraceae DrosophyllaceaeDryadaceae

Duabangaceae Duckeodendraceae Dulongiaceae Durantaceae Durionaceae Dysphani

aceae

Ebenaceae Eccremidaceae Ecdeiocoleaceae Echiaceae Echinopaceae Ehretiaceae Ela

eagnaceae ElaeocarpaceaeElegiaceae Elatinaceae Ellisiaceae Ellisiophyllaceae Elodea

ceae Elytranthaceae Embeliaceae Emblingiaceae EmmotaceaeEmpetraceae Encepha

lartaceae Engelhardtiaceae Enhalaceae Epacridaceae Ephedraceae Ephemeranthacea

eEpilobiaceae Epimediaceae Eragrostidaceae Eremolepidaceae Eremosynaceae Erica

ceae Erinaceae EriocaulaceaeEriogonaceae Eriospermaceae Erodiaceae Erycibaceae

Eryngiaceae Erysimaceae Erythroniaceae ErythropalaceaeErythrospermaceae Erythro

xylaceae Escalloniaceae Eschscholziaceae Eucomidaceae Eucommiaceae Eucryphiace

aeEuonymaceae Eupatoriaceae Euphorbiaceae Euphrasiaceae Euphroniaceae Eupo

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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matiaceae Eupteleaceae EuryalaceaeEustrephaceae Euthemidaceae Eutocaceae Exb

ucklandiaceae Exocarpaceae

Fabaceae Fagaceae Falcatifoliaceae Ferulaceae Festucaceae Ficaceae Ficoidaceae Fit

zroyaceae FlacourtiaceaeFlagellariaceae Flindersiaceae Foetidiaceae Forestieraceae

Fothergillaceae Fouquieriaceae Fragariaceae FrancoaceaeFrangulaceae Frankeniacea

e Fraxinaceae Fraxinellaceae Freycinetiaceae Fritillariaceae Fuchsiaceae Fumariacea

eFunkiaceae

Gaiadendraceae Galacaceae Galanthaceae Galaxiaceae Galedupaceae Galeniaceae G

aliaceae Garciniaceae GardeniaceaeGardneriaceae Garryaceae Geissolomataceae Ge

itonoplesiaceae Gelsemiaceae Geniostomaceae GentianaceaeGeonomataceae Geosir

idaceae Geraniaceae Gerrardinaceae Gesneriaceae Gethyllidaceae Gilliesiaceae Gina

lloaceaeGinkgoaceae Gisekiaceae Gladiolaceae Glaucidiaceae Glechomaceae Glinace

ae Globulariaceae Gnaphaliaceae GnetaceaeGoetzeaceae Gomortegaceae Gomphiac

eae Gomphrenaceae Gonystylaceae Goodeniaceae Gordoniaceae GouaniaceaeGoup

iaceae Gramineae Gratiolaceae Grewiaceae Greyiaceae Grielaceae Griseliniaceae Gr

onoviaceae GrossulariaceaeGrubbiaceae Guaiacanaceae Guamatelaceae Guettardac

eae Gunneraceae Gustaviaceae Guttiferae GyrocarpaceaeGyrostemonaceae

Hachetteaceae Haemanthaceae Haemodoraceae Halesiaceae Hallieraceae Halocarpa

ceae Halophilaceae HalophytaceaeHaloragaceae Hamamelidaceae Hanguanaceae H

aptanthaceae Harmandaceae Hebenstretiaceae HectorellaceaeHederaceae Hedyosm

aceae Hedyotidaceae Hedysaraceae Heisteriaceae Heleniaceae Heliamphoraceae H

elianthaceaeHelianthemaceae Helichrysaceae Heliconiaceae Helicteraceae Heliotropi

aceae Helleboraceae Heloniadaceae HelosidaceaeHelwingiaceae Hemerocallidaceae

Hemimeridaceae Hemiphylacaceae Henriqueziaceae Henslowiaceae Hermanniaceae

Hernandiaceae Herniariaceae Herreriaceae Hesperocallidaceae Heterantheraceae He

teropyxidaceae HeterostylaceaeHibbertiaceae Hibiscaceae Hilleriaceae Himantandra

ceae Hippocastanaceae Hippocrateaceae HippomanaceaeHippophaeaceae Hippurida

ceae Hirtellaceae Holacanthaceae Homaliaceae Hopkinsiaceae Hoplestigmataceae H

ordeaceaeHornschuchiaceae Hortensiaceae Hortoniaceae Hostaceae Houstoniaceae

Huaceae Huerteaceae HugoniaceaeHumbertiaceae Humiriaceae Hyacinthaceae Hyd

atellaceae Hydnoraceae Hydrangeaceae Hydrastidaceae HydrillaceaeHydrocharitacea

e Hydrocotylaceae Hydrogetonaceae Hydroleaceae Hydropeltidaceae Hydrophylacac

eae HydrophyllaceaeHydrostachyaceae Hymenocardiaceae Hypecoaceae Hyperanth

eraceae Hypericaceae Hypopityaceae HypoxidaceaeHypseocharitaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta

ceae ImperatoriaceaeInocarpaceae Inulaceae Ionidiaceae Iriarteaceae Iridaceae Irvi

ngiaceae Isnardiaceae Isophysidaceae IteaceaeStrasburgeriaceae Ixiaceae Ixioliriace

ae Ixonanthaceae

Jalapaceae Jamboliferaceae Japonoliriaceae Jasionaceae Jasminiaceae Johnsoniaceae

Joinvilleaceae JuglandaceaeJulianiaceae Juncaceae Juncaginaceae Juniperaceae Jus

siaeaceae Justiciaceae

Kadsuraceae Kaliphoraceae Kalmiaceae Kaniaceae Kiggelariaceae Kingdoniaceae Kin

giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae

Krameriaceae

Labiatae Lacandoniaceae Lachenaliaceae Lacistemataceae Lactoridaceae Lactucaceae

Lagerstroemiaceaee LamiaceaeLampsanaceae Lanariaceae Langsdorffiaceae Lantan

aceae Lapageriaceae Lardizabalaceae Lasiopetalaceae LauraceaeLawsoniaceae Laxm

anniaceae Lecythidaceae Ledaceae Ledocarpaceae Leeaceae Leguminosae Leitneriac

eae LemnaceaeLennoaceae Lentibulariaceae Lentiscaceae Leoniaceae Leonticaceae

Lepidobotryaceae LepidocarpaceaeLepidocaryaceae Lepidocerataceae Leptaulaceae

Leptospermaceae Lepturaceae Lepuropetalaceae LeuctenbergiaceaeLibocedraceae

Licaniaceae Ligustraceae Lilaeaceae Liliaceae Limeaceae Limnanthaceae Limnocharit

aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace

ae Linderniaceae Lindleyaceae Linnaeaceae LippayaceaeLiriaceae Liriodendraceae Li

ssocarpaceae Littorellaceae Loasaceae Lobeliaceaee Loganiaceae LomandraceaeLoni

ceraceae Lopeziaceae Lophiocarpaceae Lophiolaceae Lophiraceae Lophophytaceae L

ophopyxidaceae LoranthaceaeLotaceae Lowiaceae Lupulaceaee Luxemburgiaceae Lu

zuriagaceae Lychnidaceae Lyciaceae LyginiaceaeLygodisodeaceae Lysimachiaceae Ly

thraceae

Macarisiaceae Mackinlayaceae Madiaceae Maesaceae Magnoliaceae Malaceae Male

sherbiaceae MalortieaceaeMalpighiaceae Malvaceae Mangiaceae Manicariaceae Ma

paniaceae Marantaceae Marathraceae MarcgraviaceaeMartyniaceae Mastixiaceae

Matricariaceae Maundiaceae Mayacaceae Medeolaceae Medusagynaceae Peridiscac

eaeMelaleucaceae Melampyraceae Melanophyllaceae Melanthiaceae Melastomatac

eae Meliaceae Melianthaceae MelicaceaeMeliosmaceae Melittidaceae Melochiacea

e Memecylaceae Mendonciaceae Menispermaceae Menthaceae MenyanthaceaeMe

nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

Nypaceae Nyssaceae

Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

ceae Papilionaceae Pappophoraceae PapyraceaeParacryphiaceae Parasitaxaceae Par

ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

49

cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

50

ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 40: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

40

inghamiales Cunoniales Cupressales Cycadales Cyclanthales Cymodoceales Cynarales

CynomorialesCyperales Cyrillales Cytinales

Daphnales Daphniphyllales Dasypogonales Datiscales Degeneriales Desfontainiales D

iapensiales Didymelales DillenialesDioales Dioncophyllales Dioscoreales Diospyrales

Dipentodontales Dipsacales Droserales

Ebenales Echiales Elaeagnales Elaeocarpales Elatinales Elodeales Emmotales Empetr

ales Ephedrales Ericales EriocaulalesErythropalaless Escalloniales Eucommiales Euph

orbiales Eupomatiales Eupteleales Euryalaless

Fabales Fagales Falcatifoliales Ficales Flacourtiales Flagellariales Fouquieriales Franc

oales Frangulales

Galiales Garryales Geissolomatales Gentianales Geraniales Gesneriales Ginkgoales G

laucidiales Globulariales GnetalesGoodeniales Greyiales Griseliniales Grossulariales

Grubbiales Gunnerales Gyrocarpales Gyrostemonales

Haemodorales Haloragales Hamamelidales Hanguanales Heisteriales Helleborales He

lwingiales Himantandrales HippuridalesHomaliales Hortensiales Huales Huerteales

Hydatellales Hydnorales Hydrangeales Hydrastidales HydrocharitalesHydropeltidales

Hydrostachyales Hypericales Hypoxidales

Icacinales Illiciales Iridales Irvingales Iteales Ixiales

Jasminales Juglandales Julianiales Juncaginales Juncales

Lacistematales Lactoridales Lamiales Lardizabalales Laurales Lecythidales Ledocarpal

es Leitneriales LentibularialesLigustrales Liliales Limnanthales Linales Liriales Loasal

es Lobeliales Loganiales Lonicerales Loranthales LowialesLythrales

Magnoliales Malpighiales Malvales Marathrales Marcgraviales Mayacales Medusagy

nales Medusandrales MelanthialesMelastomatales Meliales Melianthales Meliosmal

es Menispermales Menyanthales Metteniusales MitrastemonalesMiyoshiales Moni

miales Moringales Musales Myricales Myristicales Myrothamnales Myrsinales Myrta

les

Najadales Nandinales Narcissales Nartheciales Nelumbonales Nepenthales Neuradal

es Nitrariales Nolanales NothofagalesNyctaginales Nymphaeales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

41

Ochnales Oenotherales Olacales Oleales Onagrales Oncothecales Opuntiales Orchid

ales Oxalidales

Saxifragales Pandales Pandanales Papaverales Paracryphiales Parasitaxales Paridales

Parnassiales PassifloralesPenaeales Pennantiales Pentaphragmatales Peridiscales P

etiveriales Petrosaviales Phellinales Philydrales PhyllanthalesPhysenales Phytolaccal

es Picramniales Pinales Pinguiculales Piperales Pittosporales Plantaginales Platanale

sPlumbaginales Poales Podocarpales Podophyllales Podostemales Polemoniales Pol

ygalales Polygonales PontederialesPortulacales Posidoniales Potamogetonales Primu

lales Proteales

Quercales Quillajales Quintiniales

Rafflesiales Ranunculales Rapateales Resedales Restionales Rhabdodendrales Rhamn

ales Rhinanthales RhizophoralesRhodorales Rhoipteleales Roridulales Rosales Rouss

eales Rubiales Ruppiales Rutales

Sabiales Salicales Salvadorales Samolales Samydales Sanguisorbales Santalales Sapin

dales Sapotales SarracenialesSaxegotheales Saxifragales Scheuchzeriales Sciadopitya

les Scleranthales Scrophulariales Scyphostegiales SedalesSilenales Simmondsiales S

milacales Solanales Sphenocleales Sphenostemonales Stangeriales Stellariales Stemo

nalesStilbales Stylidiales Styracales Surianales

Taccales Tamales Tamaricales Taxales Taxodiales Tecophilaeales Terebinthales Tern

stroemiales Theales TheligonalesThymelaeales Tiliales Tofieldiales Torricelliales Tov

ariales Tribelales Trilliales Trimeniales Triuridales TrochodendralesTropaeolales Turn

erales Typhales

Ulmales Urticales

Vacciniales Vahliales Vallisneriales Velloziales Veratrales Verbenales Viburnales Vinc

ales Violales Viscales VitalesVochysiales

Welwitschiales Winterales

Xanthorrhoeales Ximeniales Xyridales

Zamiales Zingiberales Zosterales Zygophyllales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

42

ALPHABETICAL LISTING OF ALL FAMILY NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | K | L | M | N | O | P | Q | R | S | T | U | V | W | X | Y

|Z

Back to Home

In some cases clicking on a family synonym will send you directly to the subfamily in

which the genus of the basionym is to be found

Abietaceae Abolbodaceae Abrophyllaceae Acalyphaceae Acanthaceae [Acanthochla

mydaceae] Acarnaceaee AceraceaeAchariaceae Achatocarpaceae Achradaceae Achy

ranthaceae Acmopylaceae Acoraceae Acristaceae ActaeaceaeActinidiaceae Actinota

ceae Amentotaxaceae Adenogrammaceae Adoxaceae Aegialitidaceae Aegicerataceae

Aegilopaceae Aeginetiaceaee Aegiphilaceae Aesculaceae Aextoxicaceae Agapanthac

eae Agavaceae AgdestidaceaeAgialidaceae Agrimoniaceae Agrostidaceae Ailanthace

ae Aitoniaceae Aizoaceae Akaniaceae Alangiaceae AlchemillaceaeAldrovandaceae A

lismataceae Alliaceae Allioniaceae Allophylaceae Aloaceae Alopecuraceae Alpiniacea

e AlseuosmiaceaeAlsinaceae Alsodeiaceae Alstroemeriaceae Altingiaceae Alzateacea

e Amaranthaceae Amaryllidaceae Amborellaceae Ambrosiaceae Amentotaxaceae A

mmanniaceae Ammiaceae Amomaceae Ampelidopsaceae Amygdalaceae Amyridacea

eAnacardiaceae Anagallidaceae Anarthriaceae Anchusaceae Ancistrocladaceae Andr

omedaceae AndropogonaceaeAndrostachyaceae Androsynaceae Anemarrhenaceae

Anemonaceae Angelicaceae Anisophylleaceae AnnonaceaeAnomochloaceae Anopter

aceae Anrederaceae Anthemidaceae Anthericaceae Anthobolaceae Antidesmataceae

Antirrhinaceae Antoniaceae Aparinaceae Aphanopetalaceae Aphloiaceae Aphyllanth

aceae Apiaceae ApocynaceaeApodanthaceae Aponogetonaceae Aporusaceae Apose

ridaceae Apostasiaceae Aptandraceae AquifoliaceaeAquilariaceae Aquilegiaceae A

raceae Aragoaceae Araliaceae Aralidiaceaee Araucariaceae Arbutaceae Arceuthidace

aeArceuthobiaceae Arctostaphylaceae Arctotidaceae Ardisiaceae Arecaceae Argoph

yllaceae Arisaraceae AristolochiaceaeAristoteliaceae Arjonaceae Armeriaceae Artem

isiaceae Arthrotaxidaceae Artocarpaceae Arundinaceae ArundinellaceaeAsaraceae A

sclepiadaceae Ascyraceaee Asiraceae Asparagaceae Asperulaceae Asphodelaceae As

pidistraceaeeAsteliaceae Asteraceae Asteranthaceae Asteropeiaceae Astragalaceae

Astrocarpaceae AthanasiaceaeAtherospermataceae Atriplicaceae Atropaceae Aucuba

ceae Aurantiaceae Austrobaileyaceae Austrotaxaceae AvenaceaeAverrhoaceae Avet

raceae Avicenniaceae Azaleaceae Azimaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

43

Balanitaceae Balanopaceae Balanophoraceae Balsameaceae Balsaminaceae Bambusa

ceae Barbaceniaceae BarbeuiaceaeBarbeyaceae Barclayaceae Barringtoniaceae Base

llaceae Bataceae Baueraceae Bauhiniaceae Baxteriaceae BegoniaceaeBehniaceae B

elangeraceae Belloniaceae Bembiciaceae Berberidaceae Berberidopsidaceae Berryac

eaee BersamaceaeBertyaceae Berzeliaceae Besleriaceae Betaceae Betulaceae Biebe

rsteiniaceae Bifariaceae Bignoniaceae BischofiaceaeBixaceae Blakeaceae Blandfordia

ceae Blattiaceae Blepharocaryaceae Blitaceae Blyxaceae Boerlagellaceae Bolivaracea

eBombacaceae Bonnetiaceae Bontiaceae Boopidaceae Boraginaceae Borassaceae B

oroniaceae BoryaceaeBotryodendraceae Bougainvilleaceae Boweniaceae Brassicacea

e Brachycaulaceae Bretschneideraceae BrexiaceaeBromeliaceae Brownlowiaceae Br

unelliaceae Bruniaceae Brunoniaceae Brunsvigiaceae Bryoniaceae BuchneraceaeBuci

daceae Buddlejaceae Buglossaceae Bulbocodiaceae Bumeliaceae Bupleuraceae Burc

hardiaceae BurmanniaceaeBurseraceae Butneriaceae Butomaceae Buxaceae Byblida

ceae Byttneriaceae

Cabombaceae Cacaoaceae Cactaceae Caesalpiniaceae Caladiaceae Calamaceae Calce

olariaceae CalectasiaceaeCalendulaceae Callaceae Callicomaceae Calligonaceae Calli

traceae Callitrichaceae Calochortaceae CalophyllaceaeCalthaceae Calycanthaceae Ca

lyceraceae Cambogiaceae Camelliaceae Campanulaceae CampynemataceaeCanacom

yricaceae Canellaceae Cannabaceae Cannaceae Canopodaceae Canotiaceae Cansjera

ceae CapparaceaeCaprariaceae Caprifoliaceae Cardiopteridaceae Carduaceae Carica

ceae Carissaceae Carlemanniaceae CarpinaceaeCarpodetaceae Cartonemataceae Ca

ryocaraceae Caryophyllaceae Casearieaceae Cassiaceae CassipoureaceaeCassythacea

e Castaneaceae Castelaceae Casuarinaceae Catesbaeaceae Cathedraceae Cecropiace

ae CedraceaeCedrelaceae Celastraceae Celosiaceae Celtidaceae Centaureaceae Cen

trolepidaceae Centroplacaceae CepaceaeCephalanthaceae Cephalotaceae Cephalota

xaceae Cerastiaceae Ceratoniaceae Ceratophyllaceae CerberaceaeCercidiphyllaceae

Cercocarpaceae Cercodiaceae Cereaceae Cerinthaceae Ceroxylaceae Cestraceae Cev

alliaceaeChailletiaceae Chamaedoraceae Chamaemoraceae Chamelauciaceae Chaun

ochitonaceae Chelidoniaceae ChelonaceaeChenopodiaceae Chimonanthaceae Chingi

thamnaceae Chionographidaceae Chiranthodendraceae ChironiaceaeChlaenaceae Ch

loanthaceae Chloranthaceae Chloridaceae Chlorogalaceae Chrysobalanaceae Cicerac

eae CichoriaceaeCimicifugaceae Cinchoniaceae Circaeaceae Circaeasteraceae Cissac

eae Cistaceae Citraceae Clematidaceae CleomaceaeClethraceae Clusiaceae Cneorac

eae Cnestidaceae Cnicaceae Cobaeaceae Cochlospermaceae Cocosaceae Coffeaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

44

Colchicaceae Coleogynaceae Columelliaceae Combretaceae Commelinaceae Comocl

adiaceae Compositae CompsoaceaeConantheraceae Connaraceae Conostylidaceae C

onvallariaceae Convolvulaceae Cordiaceae CoreopsidaceaeCoriandraceae Coriariacea

e Coridaceae Corispermaceae Cornaceae Corokiaceae Coronillaceae Correaceae Cor

rigiolaceaeCorsiaceae Corydalaceae Corylaceae Corynocarpaceae Coryphaceae Cost

aceae Cotyledonaceae CoulaceaeCoutariaceae Coutoubiaceae Crassulaceae Crescen

tiaceae Cressaceae Crinaceae Crocaceae CroomiaceaeCrossosomataceae Crotonacea

e Cruciferae Cryptaceae Crypteroniaceae Cryptocorynaceae CryptomeriaceaeCtenol

ophonaceae Cucurbitaceae Cunninghamiaceae Cunoniaceae Cupressaceae Curcumac

eae Curtisiaceae CuscutaceaeCyananthaceae Cyanastraceae Cynanchaceae Cyanella

ceae Cycadaceae Cyclanthaceae CyclantheraceaeCyclocheilaceae Cydoniaceae Cymo

doceaceae Cynaraceae Cynocrambaceae Cynomoriaceae Cyperaceae CyphiaceaeCyp

hocarpaceae Cypripediaceae Cyrillaceae Cyrtandraceae Cyrtanthaceae Cytinaceae

Dacrycarpaceae Dactylanthaceae Damasoniaceae Daphnaceae Daphniphyllaceae Das

ypogonaceae DatiscaceaeDaturaceae Daucaceae Davidiaceae Davidsoniaceae Decais

neaceae Deeringiaceae Degeneriaceae DelphiniaceaeDendrophthoaceae Columelliac

eae Detariaceae Dialypetalanthaceae Dianellaceae Dianthaceae DiapensiaceaeDicha

petalaceae Dichondraceae Diclidantheraceae Dicrastylidiaceae Dictamnaceae Didiere

aceae DidymelaceaeDidymocarpaceae Diegodendraceae Diervillaceae Digitalidaceae

e Dilatridaceae Dilleniaceae Dioaceae DionaeaceaeDioncophyllaceae Dioscoreaceae

Diosmaceae Diospyraceae Dipentodontaceae Diphylleiaceae Diplolaenaceae Dipsaca

ceaeDipterocarpaceae Dirachmaceae Disanthaceae Diselmaceae Dodonaeaceae Do

mbeyaceae Donatiaceae DorsteniaceaeDortmannaceae Doryanthaceae Drabaceae D

racaenaceae Dracontiaceae Drimyidaceae Droseraceae DrosophyllaceaeDryadaceae

Duabangaceae Duckeodendraceae Dulongiaceae Durantaceae Durionaceae Dysphani

aceae

Ebenaceae Eccremidaceae Ecdeiocoleaceae Echiaceae Echinopaceae Ehretiaceae Ela

eagnaceae ElaeocarpaceaeElegiaceae Elatinaceae Ellisiaceae Ellisiophyllaceae Elodea

ceae Elytranthaceae Embeliaceae Emblingiaceae EmmotaceaeEmpetraceae Encepha

lartaceae Engelhardtiaceae Enhalaceae Epacridaceae Ephedraceae Ephemeranthacea

eEpilobiaceae Epimediaceae Eragrostidaceae Eremolepidaceae Eremosynaceae Erica

ceae Erinaceae EriocaulaceaeEriogonaceae Eriospermaceae Erodiaceae Erycibaceae

Eryngiaceae Erysimaceae Erythroniaceae ErythropalaceaeErythrospermaceae Erythro

xylaceae Escalloniaceae Eschscholziaceae Eucomidaceae Eucommiaceae Eucryphiace

aeEuonymaceae Eupatoriaceae Euphorbiaceae Euphrasiaceae Euphroniaceae Eupo

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

45

matiaceae Eupteleaceae EuryalaceaeEustrephaceae Euthemidaceae Eutocaceae Exb

ucklandiaceae Exocarpaceae

Fabaceae Fagaceae Falcatifoliaceae Ferulaceae Festucaceae Ficaceae Ficoidaceae Fit

zroyaceae FlacourtiaceaeFlagellariaceae Flindersiaceae Foetidiaceae Forestieraceae

Fothergillaceae Fouquieriaceae Fragariaceae FrancoaceaeFrangulaceae Frankeniacea

e Fraxinaceae Fraxinellaceae Freycinetiaceae Fritillariaceae Fuchsiaceae Fumariacea

eFunkiaceae

Gaiadendraceae Galacaceae Galanthaceae Galaxiaceae Galedupaceae Galeniaceae G

aliaceae Garciniaceae GardeniaceaeGardneriaceae Garryaceae Geissolomataceae Ge

itonoplesiaceae Gelsemiaceae Geniostomaceae GentianaceaeGeonomataceae Geosir

idaceae Geraniaceae Gerrardinaceae Gesneriaceae Gethyllidaceae Gilliesiaceae Gina

lloaceaeGinkgoaceae Gisekiaceae Gladiolaceae Glaucidiaceae Glechomaceae Glinace

ae Globulariaceae Gnaphaliaceae GnetaceaeGoetzeaceae Gomortegaceae Gomphiac

eae Gomphrenaceae Gonystylaceae Goodeniaceae Gordoniaceae GouaniaceaeGoup

iaceae Gramineae Gratiolaceae Grewiaceae Greyiaceae Grielaceae Griseliniaceae Gr

onoviaceae GrossulariaceaeGrubbiaceae Guaiacanaceae Guamatelaceae Guettardac

eae Gunneraceae Gustaviaceae Guttiferae GyrocarpaceaeGyrostemonaceae

Hachetteaceae Haemanthaceae Haemodoraceae Halesiaceae Hallieraceae Halocarpa

ceae Halophilaceae HalophytaceaeHaloragaceae Hamamelidaceae Hanguanaceae H

aptanthaceae Harmandaceae Hebenstretiaceae HectorellaceaeHederaceae Hedyosm

aceae Hedyotidaceae Hedysaraceae Heisteriaceae Heleniaceae Heliamphoraceae H

elianthaceaeHelianthemaceae Helichrysaceae Heliconiaceae Helicteraceae Heliotropi

aceae Helleboraceae Heloniadaceae HelosidaceaeHelwingiaceae Hemerocallidaceae

Hemimeridaceae Hemiphylacaceae Henriqueziaceae Henslowiaceae Hermanniaceae

Hernandiaceae Herniariaceae Herreriaceae Hesperocallidaceae Heterantheraceae He

teropyxidaceae HeterostylaceaeHibbertiaceae Hibiscaceae Hilleriaceae Himantandra

ceae Hippocastanaceae Hippocrateaceae HippomanaceaeHippophaeaceae Hippurida

ceae Hirtellaceae Holacanthaceae Homaliaceae Hopkinsiaceae Hoplestigmataceae H

ordeaceaeHornschuchiaceae Hortensiaceae Hortoniaceae Hostaceae Houstoniaceae

Huaceae Huerteaceae HugoniaceaeHumbertiaceae Humiriaceae Hyacinthaceae Hyd

atellaceae Hydnoraceae Hydrangeaceae Hydrastidaceae HydrillaceaeHydrocharitacea

e Hydrocotylaceae Hydrogetonaceae Hydroleaceae Hydropeltidaceae Hydrophylacac

eae HydrophyllaceaeHydrostachyaceae Hymenocardiaceae Hypecoaceae Hyperanth

eraceae Hypericaceae Hypopityaceae HypoxidaceaeHypseocharitaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

46

Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta

ceae ImperatoriaceaeInocarpaceae Inulaceae Ionidiaceae Iriarteaceae Iridaceae Irvi

ngiaceae Isnardiaceae Isophysidaceae IteaceaeStrasburgeriaceae Ixiaceae Ixioliriace

ae Ixonanthaceae

Jalapaceae Jamboliferaceae Japonoliriaceae Jasionaceae Jasminiaceae Johnsoniaceae

Joinvilleaceae JuglandaceaeJulianiaceae Juncaceae Juncaginaceae Juniperaceae Jus

siaeaceae Justiciaceae

Kadsuraceae Kaliphoraceae Kalmiaceae Kaniaceae Kiggelariaceae Kingdoniaceae Kin

giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae

Krameriaceae

Labiatae Lacandoniaceae Lachenaliaceae Lacistemataceae Lactoridaceae Lactucaceae

Lagerstroemiaceaee LamiaceaeLampsanaceae Lanariaceae Langsdorffiaceae Lantan

aceae Lapageriaceae Lardizabalaceae Lasiopetalaceae LauraceaeLawsoniaceae Laxm

anniaceae Lecythidaceae Ledaceae Ledocarpaceae Leeaceae Leguminosae Leitneriac

eae LemnaceaeLennoaceae Lentibulariaceae Lentiscaceae Leoniaceae Leonticaceae

Lepidobotryaceae LepidocarpaceaeLepidocaryaceae Lepidocerataceae Leptaulaceae

Leptospermaceae Lepturaceae Lepuropetalaceae LeuctenbergiaceaeLibocedraceae

Licaniaceae Ligustraceae Lilaeaceae Liliaceae Limeaceae Limnanthaceae Limnocharit

aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace

ae Linderniaceae Lindleyaceae Linnaeaceae LippayaceaeLiriaceae Liriodendraceae Li

ssocarpaceae Littorellaceae Loasaceae Lobeliaceaee Loganiaceae LomandraceaeLoni

ceraceae Lopeziaceae Lophiocarpaceae Lophiolaceae Lophiraceae Lophophytaceae L

ophopyxidaceae LoranthaceaeLotaceae Lowiaceae Lupulaceaee Luxemburgiaceae Lu

zuriagaceae Lychnidaceae Lyciaceae LyginiaceaeLygodisodeaceae Lysimachiaceae Ly

thraceae

Macarisiaceae Mackinlayaceae Madiaceae Maesaceae Magnoliaceae Malaceae Male

sherbiaceae MalortieaceaeMalpighiaceae Malvaceae Mangiaceae Manicariaceae Ma

paniaceae Marantaceae Marathraceae MarcgraviaceaeMartyniaceae Mastixiaceae

Matricariaceae Maundiaceae Mayacaceae Medeolaceae Medusagynaceae Peridiscac

eaeMelaleucaceae Melampyraceae Melanophyllaceae Melanthiaceae Melastomatac

eae Meliaceae Melianthaceae MelicaceaeMeliosmaceae Melittidaceae Melochiacea

e Memecylaceae Mendonciaceae Menispermaceae Menthaceae MenyanthaceaeMe

nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

47

Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

Nypaceae Nyssaceae

Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

ceae Papilionaceae Pappophoraceae PapyraceaeParacryphiaceae Parasitaxaceae Par

ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

48

hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

49

cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

50

ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 41: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

41

Ochnales Oenotherales Olacales Oleales Onagrales Oncothecales Opuntiales Orchid

ales Oxalidales

Saxifragales Pandales Pandanales Papaverales Paracryphiales Parasitaxales Paridales

Parnassiales PassifloralesPenaeales Pennantiales Pentaphragmatales Peridiscales P

etiveriales Petrosaviales Phellinales Philydrales PhyllanthalesPhysenales Phytolaccal

es Picramniales Pinales Pinguiculales Piperales Pittosporales Plantaginales Platanale

sPlumbaginales Poales Podocarpales Podophyllales Podostemales Polemoniales Pol

ygalales Polygonales PontederialesPortulacales Posidoniales Potamogetonales Primu

lales Proteales

Quercales Quillajales Quintiniales

Rafflesiales Ranunculales Rapateales Resedales Restionales Rhabdodendrales Rhamn

ales Rhinanthales RhizophoralesRhodorales Rhoipteleales Roridulales Rosales Rouss

eales Rubiales Ruppiales Rutales

Sabiales Salicales Salvadorales Samolales Samydales Sanguisorbales Santalales Sapin

dales Sapotales SarracenialesSaxegotheales Saxifragales Scheuchzeriales Sciadopitya

les Scleranthales Scrophulariales Scyphostegiales SedalesSilenales Simmondsiales S

milacales Solanales Sphenocleales Sphenostemonales Stangeriales Stellariales Stemo

nalesStilbales Stylidiales Styracales Surianales

Taccales Tamales Tamaricales Taxales Taxodiales Tecophilaeales Terebinthales Tern

stroemiales Theales TheligonalesThymelaeales Tiliales Tofieldiales Torricelliales Tov

ariales Tribelales Trilliales Trimeniales Triuridales TrochodendralesTropaeolales Turn

erales Typhales

Ulmales Urticales

Vacciniales Vahliales Vallisneriales Velloziales Veratrales Verbenales Viburnales Vinc

ales Violales Viscales VitalesVochysiales

Welwitschiales Winterales

Xanthorrhoeales Ximeniales Xyridales

Zamiales Zingiberales Zosterales Zygophyllales

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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ALPHABETICAL LISTING OF ALL FAMILY NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | K | L | M | N | O | P | Q | R | S | T | U | V | W | X | Y

|Z

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In some cases clicking on a family synonym will send you directly to the subfamily in

which the genus of the basionym is to be found

Abietaceae Abolbodaceae Abrophyllaceae Acalyphaceae Acanthaceae [Acanthochla

mydaceae] Acarnaceaee AceraceaeAchariaceae Achatocarpaceae Achradaceae Achy

ranthaceae Acmopylaceae Acoraceae Acristaceae ActaeaceaeActinidiaceae Actinota

ceae Amentotaxaceae Adenogrammaceae Adoxaceae Aegialitidaceae Aegicerataceae

Aegilopaceae Aeginetiaceaee Aegiphilaceae Aesculaceae Aextoxicaceae Agapanthac

eae Agavaceae AgdestidaceaeAgialidaceae Agrimoniaceae Agrostidaceae Ailanthace

ae Aitoniaceae Aizoaceae Akaniaceae Alangiaceae AlchemillaceaeAldrovandaceae A

lismataceae Alliaceae Allioniaceae Allophylaceae Aloaceae Alopecuraceae Alpiniacea

e AlseuosmiaceaeAlsinaceae Alsodeiaceae Alstroemeriaceae Altingiaceae Alzateacea

e Amaranthaceae Amaryllidaceae Amborellaceae Ambrosiaceae Amentotaxaceae A

mmanniaceae Ammiaceae Amomaceae Ampelidopsaceae Amygdalaceae Amyridacea

eAnacardiaceae Anagallidaceae Anarthriaceae Anchusaceae Ancistrocladaceae Andr

omedaceae AndropogonaceaeAndrostachyaceae Androsynaceae Anemarrhenaceae

Anemonaceae Angelicaceae Anisophylleaceae AnnonaceaeAnomochloaceae Anopter

aceae Anrederaceae Anthemidaceae Anthericaceae Anthobolaceae Antidesmataceae

Antirrhinaceae Antoniaceae Aparinaceae Aphanopetalaceae Aphloiaceae Aphyllanth

aceae Apiaceae ApocynaceaeApodanthaceae Aponogetonaceae Aporusaceae Apose

ridaceae Apostasiaceae Aptandraceae AquifoliaceaeAquilariaceae Aquilegiaceae A

raceae Aragoaceae Araliaceae Aralidiaceaee Araucariaceae Arbutaceae Arceuthidace

aeArceuthobiaceae Arctostaphylaceae Arctotidaceae Ardisiaceae Arecaceae Argoph

yllaceae Arisaraceae AristolochiaceaeAristoteliaceae Arjonaceae Armeriaceae Artem

isiaceae Arthrotaxidaceae Artocarpaceae Arundinaceae ArundinellaceaeAsaraceae A

sclepiadaceae Ascyraceaee Asiraceae Asparagaceae Asperulaceae Asphodelaceae As

pidistraceaeeAsteliaceae Asteraceae Asteranthaceae Asteropeiaceae Astragalaceae

Astrocarpaceae AthanasiaceaeAtherospermataceae Atriplicaceae Atropaceae Aucuba

ceae Aurantiaceae Austrobaileyaceae Austrotaxaceae AvenaceaeAverrhoaceae Avet

raceae Avicenniaceae Azaleaceae Azimaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Balanitaceae Balanopaceae Balanophoraceae Balsameaceae Balsaminaceae Bambusa

ceae Barbaceniaceae BarbeuiaceaeBarbeyaceae Barclayaceae Barringtoniaceae Base

llaceae Bataceae Baueraceae Bauhiniaceae Baxteriaceae BegoniaceaeBehniaceae B

elangeraceae Belloniaceae Bembiciaceae Berberidaceae Berberidopsidaceae Berryac

eaee BersamaceaeBertyaceae Berzeliaceae Besleriaceae Betaceae Betulaceae Biebe

rsteiniaceae Bifariaceae Bignoniaceae BischofiaceaeBixaceae Blakeaceae Blandfordia

ceae Blattiaceae Blepharocaryaceae Blitaceae Blyxaceae Boerlagellaceae Bolivaracea

eBombacaceae Bonnetiaceae Bontiaceae Boopidaceae Boraginaceae Borassaceae B

oroniaceae BoryaceaeBotryodendraceae Bougainvilleaceae Boweniaceae Brassicacea

e Brachycaulaceae Bretschneideraceae BrexiaceaeBromeliaceae Brownlowiaceae Br

unelliaceae Bruniaceae Brunoniaceae Brunsvigiaceae Bryoniaceae BuchneraceaeBuci

daceae Buddlejaceae Buglossaceae Bulbocodiaceae Bumeliaceae Bupleuraceae Burc

hardiaceae BurmanniaceaeBurseraceae Butneriaceae Butomaceae Buxaceae Byblida

ceae Byttneriaceae

Cabombaceae Cacaoaceae Cactaceae Caesalpiniaceae Caladiaceae Calamaceae Calce

olariaceae CalectasiaceaeCalendulaceae Callaceae Callicomaceae Calligonaceae Calli

traceae Callitrichaceae Calochortaceae CalophyllaceaeCalthaceae Calycanthaceae Ca

lyceraceae Cambogiaceae Camelliaceae Campanulaceae CampynemataceaeCanacom

yricaceae Canellaceae Cannabaceae Cannaceae Canopodaceae Canotiaceae Cansjera

ceae CapparaceaeCaprariaceae Caprifoliaceae Cardiopteridaceae Carduaceae Carica

ceae Carissaceae Carlemanniaceae CarpinaceaeCarpodetaceae Cartonemataceae Ca

ryocaraceae Caryophyllaceae Casearieaceae Cassiaceae CassipoureaceaeCassythacea

e Castaneaceae Castelaceae Casuarinaceae Catesbaeaceae Cathedraceae Cecropiace

ae CedraceaeCedrelaceae Celastraceae Celosiaceae Celtidaceae Centaureaceae Cen

trolepidaceae Centroplacaceae CepaceaeCephalanthaceae Cephalotaceae Cephalota

xaceae Cerastiaceae Ceratoniaceae Ceratophyllaceae CerberaceaeCercidiphyllaceae

Cercocarpaceae Cercodiaceae Cereaceae Cerinthaceae Ceroxylaceae Cestraceae Cev

alliaceaeChailletiaceae Chamaedoraceae Chamaemoraceae Chamelauciaceae Chaun

ochitonaceae Chelidoniaceae ChelonaceaeChenopodiaceae Chimonanthaceae Chingi

thamnaceae Chionographidaceae Chiranthodendraceae ChironiaceaeChlaenaceae Ch

loanthaceae Chloranthaceae Chloridaceae Chlorogalaceae Chrysobalanaceae Cicerac

eae CichoriaceaeCimicifugaceae Cinchoniaceae Circaeaceae Circaeasteraceae Cissac

eae Cistaceae Citraceae Clematidaceae CleomaceaeClethraceae Clusiaceae Cneorac

eae Cnestidaceae Cnicaceae Cobaeaceae Cochlospermaceae Cocosaceae Coffeaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Colchicaceae Coleogynaceae Columelliaceae Combretaceae Commelinaceae Comocl

adiaceae Compositae CompsoaceaeConantheraceae Connaraceae Conostylidaceae C

onvallariaceae Convolvulaceae Cordiaceae CoreopsidaceaeCoriandraceae Coriariacea

e Coridaceae Corispermaceae Cornaceae Corokiaceae Coronillaceae Correaceae Cor

rigiolaceaeCorsiaceae Corydalaceae Corylaceae Corynocarpaceae Coryphaceae Cost

aceae Cotyledonaceae CoulaceaeCoutariaceae Coutoubiaceae Crassulaceae Crescen

tiaceae Cressaceae Crinaceae Crocaceae CroomiaceaeCrossosomataceae Crotonacea

e Cruciferae Cryptaceae Crypteroniaceae Cryptocorynaceae CryptomeriaceaeCtenol

ophonaceae Cucurbitaceae Cunninghamiaceae Cunoniaceae Cupressaceae Curcumac

eae Curtisiaceae CuscutaceaeCyananthaceae Cyanastraceae Cynanchaceae Cyanella

ceae Cycadaceae Cyclanthaceae CyclantheraceaeCyclocheilaceae Cydoniaceae Cymo

doceaceae Cynaraceae Cynocrambaceae Cynomoriaceae Cyperaceae CyphiaceaeCyp

hocarpaceae Cypripediaceae Cyrillaceae Cyrtandraceae Cyrtanthaceae Cytinaceae

Dacrycarpaceae Dactylanthaceae Damasoniaceae Daphnaceae Daphniphyllaceae Das

ypogonaceae DatiscaceaeDaturaceae Daucaceae Davidiaceae Davidsoniaceae Decais

neaceae Deeringiaceae Degeneriaceae DelphiniaceaeDendrophthoaceae Columelliac

eae Detariaceae Dialypetalanthaceae Dianellaceae Dianthaceae DiapensiaceaeDicha

petalaceae Dichondraceae Diclidantheraceae Dicrastylidiaceae Dictamnaceae Didiere

aceae DidymelaceaeDidymocarpaceae Diegodendraceae Diervillaceae Digitalidaceae

e Dilatridaceae Dilleniaceae Dioaceae DionaeaceaeDioncophyllaceae Dioscoreaceae

Diosmaceae Diospyraceae Dipentodontaceae Diphylleiaceae Diplolaenaceae Dipsaca

ceaeDipterocarpaceae Dirachmaceae Disanthaceae Diselmaceae Dodonaeaceae Do

mbeyaceae Donatiaceae DorsteniaceaeDortmannaceae Doryanthaceae Drabaceae D

racaenaceae Dracontiaceae Drimyidaceae Droseraceae DrosophyllaceaeDryadaceae

Duabangaceae Duckeodendraceae Dulongiaceae Durantaceae Durionaceae Dysphani

aceae

Ebenaceae Eccremidaceae Ecdeiocoleaceae Echiaceae Echinopaceae Ehretiaceae Ela

eagnaceae ElaeocarpaceaeElegiaceae Elatinaceae Ellisiaceae Ellisiophyllaceae Elodea

ceae Elytranthaceae Embeliaceae Emblingiaceae EmmotaceaeEmpetraceae Encepha

lartaceae Engelhardtiaceae Enhalaceae Epacridaceae Ephedraceae Ephemeranthacea

eEpilobiaceae Epimediaceae Eragrostidaceae Eremolepidaceae Eremosynaceae Erica

ceae Erinaceae EriocaulaceaeEriogonaceae Eriospermaceae Erodiaceae Erycibaceae

Eryngiaceae Erysimaceae Erythroniaceae ErythropalaceaeErythrospermaceae Erythro

xylaceae Escalloniaceae Eschscholziaceae Eucomidaceae Eucommiaceae Eucryphiace

aeEuonymaceae Eupatoriaceae Euphorbiaceae Euphrasiaceae Euphroniaceae Eupo

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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matiaceae Eupteleaceae EuryalaceaeEustrephaceae Euthemidaceae Eutocaceae Exb

ucklandiaceae Exocarpaceae

Fabaceae Fagaceae Falcatifoliaceae Ferulaceae Festucaceae Ficaceae Ficoidaceae Fit

zroyaceae FlacourtiaceaeFlagellariaceae Flindersiaceae Foetidiaceae Forestieraceae

Fothergillaceae Fouquieriaceae Fragariaceae FrancoaceaeFrangulaceae Frankeniacea

e Fraxinaceae Fraxinellaceae Freycinetiaceae Fritillariaceae Fuchsiaceae Fumariacea

eFunkiaceae

Gaiadendraceae Galacaceae Galanthaceae Galaxiaceae Galedupaceae Galeniaceae G

aliaceae Garciniaceae GardeniaceaeGardneriaceae Garryaceae Geissolomataceae Ge

itonoplesiaceae Gelsemiaceae Geniostomaceae GentianaceaeGeonomataceae Geosir

idaceae Geraniaceae Gerrardinaceae Gesneriaceae Gethyllidaceae Gilliesiaceae Gina

lloaceaeGinkgoaceae Gisekiaceae Gladiolaceae Glaucidiaceae Glechomaceae Glinace

ae Globulariaceae Gnaphaliaceae GnetaceaeGoetzeaceae Gomortegaceae Gomphiac

eae Gomphrenaceae Gonystylaceae Goodeniaceae Gordoniaceae GouaniaceaeGoup

iaceae Gramineae Gratiolaceae Grewiaceae Greyiaceae Grielaceae Griseliniaceae Gr

onoviaceae GrossulariaceaeGrubbiaceae Guaiacanaceae Guamatelaceae Guettardac

eae Gunneraceae Gustaviaceae Guttiferae GyrocarpaceaeGyrostemonaceae

Hachetteaceae Haemanthaceae Haemodoraceae Halesiaceae Hallieraceae Halocarpa

ceae Halophilaceae HalophytaceaeHaloragaceae Hamamelidaceae Hanguanaceae H

aptanthaceae Harmandaceae Hebenstretiaceae HectorellaceaeHederaceae Hedyosm

aceae Hedyotidaceae Hedysaraceae Heisteriaceae Heleniaceae Heliamphoraceae H

elianthaceaeHelianthemaceae Helichrysaceae Heliconiaceae Helicteraceae Heliotropi

aceae Helleboraceae Heloniadaceae HelosidaceaeHelwingiaceae Hemerocallidaceae

Hemimeridaceae Hemiphylacaceae Henriqueziaceae Henslowiaceae Hermanniaceae

Hernandiaceae Herniariaceae Herreriaceae Hesperocallidaceae Heterantheraceae He

teropyxidaceae HeterostylaceaeHibbertiaceae Hibiscaceae Hilleriaceae Himantandra

ceae Hippocastanaceae Hippocrateaceae HippomanaceaeHippophaeaceae Hippurida

ceae Hirtellaceae Holacanthaceae Homaliaceae Hopkinsiaceae Hoplestigmataceae H

ordeaceaeHornschuchiaceae Hortensiaceae Hortoniaceae Hostaceae Houstoniaceae

Huaceae Huerteaceae HugoniaceaeHumbertiaceae Humiriaceae Hyacinthaceae Hyd

atellaceae Hydnoraceae Hydrangeaceae Hydrastidaceae HydrillaceaeHydrocharitacea

e Hydrocotylaceae Hydrogetonaceae Hydroleaceae Hydropeltidaceae Hydrophylacac

eae HydrophyllaceaeHydrostachyaceae Hymenocardiaceae Hypecoaceae Hyperanth

eraceae Hypericaceae Hypopityaceae HypoxidaceaeHypseocharitaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta

ceae ImperatoriaceaeInocarpaceae Inulaceae Ionidiaceae Iriarteaceae Iridaceae Irvi

ngiaceae Isnardiaceae Isophysidaceae IteaceaeStrasburgeriaceae Ixiaceae Ixioliriace

ae Ixonanthaceae

Jalapaceae Jamboliferaceae Japonoliriaceae Jasionaceae Jasminiaceae Johnsoniaceae

Joinvilleaceae JuglandaceaeJulianiaceae Juncaceae Juncaginaceae Juniperaceae Jus

siaeaceae Justiciaceae

Kadsuraceae Kaliphoraceae Kalmiaceae Kaniaceae Kiggelariaceae Kingdoniaceae Kin

giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae

Krameriaceae

Labiatae Lacandoniaceae Lachenaliaceae Lacistemataceae Lactoridaceae Lactucaceae

Lagerstroemiaceaee LamiaceaeLampsanaceae Lanariaceae Langsdorffiaceae Lantan

aceae Lapageriaceae Lardizabalaceae Lasiopetalaceae LauraceaeLawsoniaceae Laxm

anniaceae Lecythidaceae Ledaceae Ledocarpaceae Leeaceae Leguminosae Leitneriac

eae LemnaceaeLennoaceae Lentibulariaceae Lentiscaceae Leoniaceae Leonticaceae

Lepidobotryaceae LepidocarpaceaeLepidocaryaceae Lepidocerataceae Leptaulaceae

Leptospermaceae Lepturaceae Lepuropetalaceae LeuctenbergiaceaeLibocedraceae

Licaniaceae Ligustraceae Lilaeaceae Liliaceae Limeaceae Limnanthaceae Limnocharit

aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace

ae Linderniaceae Lindleyaceae Linnaeaceae LippayaceaeLiriaceae Liriodendraceae Li

ssocarpaceae Littorellaceae Loasaceae Lobeliaceaee Loganiaceae LomandraceaeLoni

ceraceae Lopeziaceae Lophiocarpaceae Lophiolaceae Lophiraceae Lophophytaceae L

ophopyxidaceae LoranthaceaeLotaceae Lowiaceae Lupulaceaee Luxemburgiaceae Lu

zuriagaceae Lychnidaceae Lyciaceae LyginiaceaeLygodisodeaceae Lysimachiaceae Ly

thraceae

Macarisiaceae Mackinlayaceae Madiaceae Maesaceae Magnoliaceae Malaceae Male

sherbiaceae MalortieaceaeMalpighiaceae Malvaceae Mangiaceae Manicariaceae Ma

paniaceae Marantaceae Marathraceae MarcgraviaceaeMartyniaceae Mastixiaceae

Matricariaceae Maundiaceae Mayacaceae Medeolaceae Medusagynaceae Peridiscac

eaeMelaleucaceae Melampyraceae Melanophyllaceae Melanthiaceae Melastomatac

eae Meliaceae Melianthaceae MelicaceaeMeliosmaceae Melittidaceae Melochiacea

e Memecylaceae Mendonciaceae Menispermaceae Menthaceae MenyanthaceaeMe

nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

Nypaceae Nyssaceae

Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

ceae Papilionaceae Pappophoraceae PapyraceaeParacryphiaceae Parasitaxaceae Par

ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

48

hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

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ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 42: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

42

ALPHABETICAL LISTING OF ALL FAMILY NAMES OF SEED PLANTS WITH LINKS

A | B | C | D | E | F | G | H | I | J | K | L | M | N | O | P | Q | R | S | T | U | V | W | X | Y

|Z

Back to Home

In some cases clicking on a family synonym will send you directly to the subfamily in

which the genus of the basionym is to be found

Abietaceae Abolbodaceae Abrophyllaceae Acalyphaceae Acanthaceae [Acanthochla

mydaceae] Acarnaceaee AceraceaeAchariaceae Achatocarpaceae Achradaceae Achy

ranthaceae Acmopylaceae Acoraceae Acristaceae ActaeaceaeActinidiaceae Actinota

ceae Amentotaxaceae Adenogrammaceae Adoxaceae Aegialitidaceae Aegicerataceae

Aegilopaceae Aeginetiaceaee Aegiphilaceae Aesculaceae Aextoxicaceae Agapanthac

eae Agavaceae AgdestidaceaeAgialidaceae Agrimoniaceae Agrostidaceae Ailanthace

ae Aitoniaceae Aizoaceae Akaniaceae Alangiaceae AlchemillaceaeAldrovandaceae A

lismataceae Alliaceae Allioniaceae Allophylaceae Aloaceae Alopecuraceae Alpiniacea

e AlseuosmiaceaeAlsinaceae Alsodeiaceae Alstroemeriaceae Altingiaceae Alzateacea

e Amaranthaceae Amaryllidaceae Amborellaceae Ambrosiaceae Amentotaxaceae A

mmanniaceae Ammiaceae Amomaceae Ampelidopsaceae Amygdalaceae Amyridacea

eAnacardiaceae Anagallidaceae Anarthriaceae Anchusaceae Ancistrocladaceae Andr

omedaceae AndropogonaceaeAndrostachyaceae Androsynaceae Anemarrhenaceae

Anemonaceae Angelicaceae Anisophylleaceae AnnonaceaeAnomochloaceae Anopter

aceae Anrederaceae Anthemidaceae Anthericaceae Anthobolaceae Antidesmataceae

Antirrhinaceae Antoniaceae Aparinaceae Aphanopetalaceae Aphloiaceae Aphyllanth

aceae Apiaceae ApocynaceaeApodanthaceae Aponogetonaceae Aporusaceae Apose

ridaceae Apostasiaceae Aptandraceae AquifoliaceaeAquilariaceae Aquilegiaceae A

raceae Aragoaceae Araliaceae Aralidiaceaee Araucariaceae Arbutaceae Arceuthidace

aeArceuthobiaceae Arctostaphylaceae Arctotidaceae Ardisiaceae Arecaceae Argoph

yllaceae Arisaraceae AristolochiaceaeAristoteliaceae Arjonaceae Armeriaceae Artem

isiaceae Arthrotaxidaceae Artocarpaceae Arundinaceae ArundinellaceaeAsaraceae A

sclepiadaceae Ascyraceaee Asiraceae Asparagaceae Asperulaceae Asphodelaceae As

pidistraceaeeAsteliaceae Asteraceae Asteranthaceae Asteropeiaceae Astragalaceae

Astrocarpaceae AthanasiaceaeAtherospermataceae Atriplicaceae Atropaceae Aucuba

ceae Aurantiaceae Austrobaileyaceae Austrotaxaceae AvenaceaeAverrhoaceae Avet

raceae Avicenniaceae Azaleaceae Azimaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

43

Balanitaceae Balanopaceae Balanophoraceae Balsameaceae Balsaminaceae Bambusa

ceae Barbaceniaceae BarbeuiaceaeBarbeyaceae Barclayaceae Barringtoniaceae Base

llaceae Bataceae Baueraceae Bauhiniaceae Baxteriaceae BegoniaceaeBehniaceae B

elangeraceae Belloniaceae Bembiciaceae Berberidaceae Berberidopsidaceae Berryac

eaee BersamaceaeBertyaceae Berzeliaceae Besleriaceae Betaceae Betulaceae Biebe

rsteiniaceae Bifariaceae Bignoniaceae BischofiaceaeBixaceae Blakeaceae Blandfordia

ceae Blattiaceae Blepharocaryaceae Blitaceae Blyxaceae Boerlagellaceae Bolivaracea

eBombacaceae Bonnetiaceae Bontiaceae Boopidaceae Boraginaceae Borassaceae B

oroniaceae BoryaceaeBotryodendraceae Bougainvilleaceae Boweniaceae Brassicacea

e Brachycaulaceae Bretschneideraceae BrexiaceaeBromeliaceae Brownlowiaceae Br

unelliaceae Bruniaceae Brunoniaceae Brunsvigiaceae Bryoniaceae BuchneraceaeBuci

daceae Buddlejaceae Buglossaceae Bulbocodiaceae Bumeliaceae Bupleuraceae Burc

hardiaceae BurmanniaceaeBurseraceae Butneriaceae Butomaceae Buxaceae Byblida

ceae Byttneriaceae

Cabombaceae Cacaoaceae Cactaceae Caesalpiniaceae Caladiaceae Calamaceae Calce

olariaceae CalectasiaceaeCalendulaceae Callaceae Callicomaceae Calligonaceae Calli

traceae Callitrichaceae Calochortaceae CalophyllaceaeCalthaceae Calycanthaceae Ca

lyceraceae Cambogiaceae Camelliaceae Campanulaceae CampynemataceaeCanacom

yricaceae Canellaceae Cannabaceae Cannaceae Canopodaceae Canotiaceae Cansjera

ceae CapparaceaeCaprariaceae Caprifoliaceae Cardiopteridaceae Carduaceae Carica

ceae Carissaceae Carlemanniaceae CarpinaceaeCarpodetaceae Cartonemataceae Ca

ryocaraceae Caryophyllaceae Casearieaceae Cassiaceae CassipoureaceaeCassythacea

e Castaneaceae Castelaceae Casuarinaceae Catesbaeaceae Cathedraceae Cecropiace

ae CedraceaeCedrelaceae Celastraceae Celosiaceae Celtidaceae Centaureaceae Cen

trolepidaceae Centroplacaceae CepaceaeCephalanthaceae Cephalotaceae Cephalota

xaceae Cerastiaceae Ceratoniaceae Ceratophyllaceae CerberaceaeCercidiphyllaceae

Cercocarpaceae Cercodiaceae Cereaceae Cerinthaceae Ceroxylaceae Cestraceae Cev

alliaceaeChailletiaceae Chamaedoraceae Chamaemoraceae Chamelauciaceae Chaun

ochitonaceae Chelidoniaceae ChelonaceaeChenopodiaceae Chimonanthaceae Chingi

thamnaceae Chionographidaceae Chiranthodendraceae ChironiaceaeChlaenaceae Ch

loanthaceae Chloranthaceae Chloridaceae Chlorogalaceae Chrysobalanaceae Cicerac

eae CichoriaceaeCimicifugaceae Cinchoniaceae Circaeaceae Circaeasteraceae Cissac

eae Cistaceae Citraceae Clematidaceae CleomaceaeClethraceae Clusiaceae Cneorac

eae Cnestidaceae Cnicaceae Cobaeaceae Cochlospermaceae Cocosaceae Coffeaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

44

Colchicaceae Coleogynaceae Columelliaceae Combretaceae Commelinaceae Comocl

adiaceae Compositae CompsoaceaeConantheraceae Connaraceae Conostylidaceae C

onvallariaceae Convolvulaceae Cordiaceae CoreopsidaceaeCoriandraceae Coriariacea

e Coridaceae Corispermaceae Cornaceae Corokiaceae Coronillaceae Correaceae Cor

rigiolaceaeCorsiaceae Corydalaceae Corylaceae Corynocarpaceae Coryphaceae Cost

aceae Cotyledonaceae CoulaceaeCoutariaceae Coutoubiaceae Crassulaceae Crescen

tiaceae Cressaceae Crinaceae Crocaceae CroomiaceaeCrossosomataceae Crotonacea

e Cruciferae Cryptaceae Crypteroniaceae Cryptocorynaceae CryptomeriaceaeCtenol

ophonaceae Cucurbitaceae Cunninghamiaceae Cunoniaceae Cupressaceae Curcumac

eae Curtisiaceae CuscutaceaeCyananthaceae Cyanastraceae Cynanchaceae Cyanella

ceae Cycadaceae Cyclanthaceae CyclantheraceaeCyclocheilaceae Cydoniaceae Cymo

doceaceae Cynaraceae Cynocrambaceae Cynomoriaceae Cyperaceae CyphiaceaeCyp

hocarpaceae Cypripediaceae Cyrillaceae Cyrtandraceae Cyrtanthaceae Cytinaceae

Dacrycarpaceae Dactylanthaceae Damasoniaceae Daphnaceae Daphniphyllaceae Das

ypogonaceae DatiscaceaeDaturaceae Daucaceae Davidiaceae Davidsoniaceae Decais

neaceae Deeringiaceae Degeneriaceae DelphiniaceaeDendrophthoaceae Columelliac

eae Detariaceae Dialypetalanthaceae Dianellaceae Dianthaceae DiapensiaceaeDicha

petalaceae Dichondraceae Diclidantheraceae Dicrastylidiaceae Dictamnaceae Didiere

aceae DidymelaceaeDidymocarpaceae Diegodendraceae Diervillaceae Digitalidaceae

e Dilatridaceae Dilleniaceae Dioaceae DionaeaceaeDioncophyllaceae Dioscoreaceae

Diosmaceae Diospyraceae Dipentodontaceae Diphylleiaceae Diplolaenaceae Dipsaca

ceaeDipterocarpaceae Dirachmaceae Disanthaceae Diselmaceae Dodonaeaceae Do

mbeyaceae Donatiaceae DorsteniaceaeDortmannaceae Doryanthaceae Drabaceae D

racaenaceae Dracontiaceae Drimyidaceae Droseraceae DrosophyllaceaeDryadaceae

Duabangaceae Duckeodendraceae Dulongiaceae Durantaceae Durionaceae Dysphani

aceae

Ebenaceae Eccremidaceae Ecdeiocoleaceae Echiaceae Echinopaceae Ehretiaceae Ela

eagnaceae ElaeocarpaceaeElegiaceae Elatinaceae Ellisiaceae Ellisiophyllaceae Elodea

ceae Elytranthaceae Embeliaceae Emblingiaceae EmmotaceaeEmpetraceae Encepha

lartaceae Engelhardtiaceae Enhalaceae Epacridaceae Ephedraceae Ephemeranthacea

eEpilobiaceae Epimediaceae Eragrostidaceae Eremolepidaceae Eremosynaceae Erica

ceae Erinaceae EriocaulaceaeEriogonaceae Eriospermaceae Erodiaceae Erycibaceae

Eryngiaceae Erysimaceae Erythroniaceae ErythropalaceaeErythrospermaceae Erythro

xylaceae Escalloniaceae Eschscholziaceae Eucomidaceae Eucommiaceae Eucryphiace

aeEuonymaceae Eupatoriaceae Euphorbiaceae Euphrasiaceae Euphroniaceae Eupo

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

45

matiaceae Eupteleaceae EuryalaceaeEustrephaceae Euthemidaceae Eutocaceae Exb

ucklandiaceae Exocarpaceae

Fabaceae Fagaceae Falcatifoliaceae Ferulaceae Festucaceae Ficaceae Ficoidaceae Fit

zroyaceae FlacourtiaceaeFlagellariaceae Flindersiaceae Foetidiaceae Forestieraceae

Fothergillaceae Fouquieriaceae Fragariaceae FrancoaceaeFrangulaceae Frankeniacea

e Fraxinaceae Fraxinellaceae Freycinetiaceae Fritillariaceae Fuchsiaceae Fumariacea

eFunkiaceae

Gaiadendraceae Galacaceae Galanthaceae Galaxiaceae Galedupaceae Galeniaceae G

aliaceae Garciniaceae GardeniaceaeGardneriaceae Garryaceae Geissolomataceae Ge

itonoplesiaceae Gelsemiaceae Geniostomaceae GentianaceaeGeonomataceae Geosir

idaceae Geraniaceae Gerrardinaceae Gesneriaceae Gethyllidaceae Gilliesiaceae Gina

lloaceaeGinkgoaceae Gisekiaceae Gladiolaceae Glaucidiaceae Glechomaceae Glinace

ae Globulariaceae Gnaphaliaceae GnetaceaeGoetzeaceae Gomortegaceae Gomphiac

eae Gomphrenaceae Gonystylaceae Goodeniaceae Gordoniaceae GouaniaceaeGoup

iaceae Gramineae Gratiolaceae Grewiaceae Greyiaceae Grielaceae Griseliniaceae Gr

onoviaceae GrossulariaceaeGrubbiaceae Guaiacanaceae Guamatelaceae Guettardac

eae Gunneraceae Gustaviaceae Guttiferae GyrocarpaceaeGyrostemonaceae

Hachetteaceae Haemanthaceae Haemodoraceae Halesiaceae Hallieraceae Halocarpa

ceae Halophilaceae HalophytaceaeHaloragaceae Hamamelidaceae Hanguanaceae H

aptanthaceae Harmandaceae Hebenstretiaceae HectorellaceaeHederaceae Hedyosm

aceae Hedyotidaceae Hedysaraceae Heisteriaceae Heleniaceae Heliamphoraceae H

elianthaceaeHelianthemaceae Helichrysaceae Heliconiaceae Helicteraceae Heliotropi

aceae Helleboraceae Heloniadaceae HelosidaceaeHelwingiaceae Hemerocallidaceae

Hemimeridaceae Hemiphylacaceae Henriqueziaceae Henslowiaceae Hermanniaceae

Hernandiaceae Herniariaceae Herreriaceae Hesperocallidaceae Heterantheraceae He

teropyxidaceae HeterostylaceaeHibbertiaceae Hibiscaceae Hilleriaceae Himantandra

ceae Hippocastanaceae Hippocrateaceae HippomanaceaeHippophaeaceae Hippurida

ceae Hirtellaceae Holacanthaceae Homaliaceae Hopkinsiaceae Hoplestigmataceae H

ordeaceaeHornschuchiaceae Hortensiaceae Hortoniaceae Hostaceae Houstoniaceae

Huaceae Huerteaceae HugoniaceaeHumbertiaceae Humiriaceae Hyacinthaceae Hyd

atellaceae Hydnoraceae Hydrangeaceae Hydrastidaceae HydrillaceaeHydrocharitacea

e Hydrocotylaceae Hydrogetonaceae Hydroleaceae Hydropeltidaceae Hydrophylacac

eae HydrophyllaceaeHydrostachyaceae Hymenocardiaceae Hypecoaceae Hyperanth

eraceae Hypericaceae Hypopityaceae HypoxidaceaeHypseocharitaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

46

Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta

ceae ImperatoriaceaeInocarpaceae Inulaceae Ionidiaceae Iriarteaceae Iridaceae Irvi

ngiaceae Isnardiaceae Isophysidaceae IteaceaeStrasburgeriaceae Ixiaceae Ixioliriace

ae Ixonanthaceae

Jalapaceae Jamboliferaceae Japonoliriaceae Jasionaceae Jasminiaceae Johnsoniaceae

Joinvilleaceae JuglandaceaeJulianiaceae Juncaceae Juncaginaceae Juniperaceae Jus

siaeaceae Justiciaceae

Kadsuraceae Kaliphoraceae Kalmiaceae Kaniaceae Kiggelariaceae Kingdoniaceae Kin

giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae

Krameriaceae

Labiatae Lacandoniaceae Lachenaliaceae Lacistemataceae Lactoridaceae Lactucaceae

Lagerstroemiaceaee LamiaceaeLampsanaceae Lanariaceae Langsdorffiaceae Lantan

aceae Lapageriaceae Lardizabalaceae Lasiopetalaceae LauraceaeLawsoniaceae Laxm

anniaceae Lecythidaceae Ledaceae Ledocarpaceae Leeaceae Leguminosae Leitneriac

eae LemnaceaeLennoaceae Lentibulariaceae Lentiscaceae Leoniaceae Leonticaceae

Lepidobotryaceae LepidocarpaceaeLepidocaryaceae Lepidocerataceae Leptaulaceae

Leptospermaceae Lepturaceae Lepuropetalaceae LeuctenbergiaceaeLibocedraceae

Licaniaceae Ligustraceae Lilaeaceae Liliaceae Limeaceae Limnanthaceae Limnocharit

aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace

ae Linderniaceae Lindleyaceae Linnaeaceae LippayaceaeLiriaceae Liriodendraceae Li

ssocarpaceae Littorellaceae Loasaceae Lobeliaceaee Loganiaceae LomandraceaeLoni

ceraceae Lopeziaceae Lophiocarpaceae Lophiolaceae Lophiraceae Lophophytaceae L

ophopyxidaceae LoranthaceaeLotaceae Lowiaceae Lupulaceaee Luxemburgiaceae Lu

zuriagaceae Lychnidaceae Lyciaceae LyginiaceaeLygodisodeaceae Lysimachiaceae Ly

thraceae

Macarisiaceae Mackinlayaceae Madiaceae Maesaceae Magnoliaceae Malaceae Male

sherbiaceae MalortieaceaeMalpighiaceae Malvaceae Mangiaceae Manicariaceae Ma

paniaceae Marantaceae Marathraceae MarcgraviaceaeMartyniaceae Mastixiaceae

Matricariaceae Maundiaceae Mayacaceae Medeolaceae Medusagynaceae Peridiscac

eaeMelaleucaceae Melampyraceae Melanophyllaceae Melanthiaceae Melastomatac

eae Meliaceae Melianthaceae MelicaceaeMeliosmaceae Melittidaceae Melochiacea

e Memecylaceae Mendonciaceae Menispermaceae Menthaceae MenyanthaceaeMe

nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

47

Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

Nypaceae Nyssaceae

Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

ceae Papilionaceae Pappophoraceae PapyraceaeParacryphiaceae Parasitaxaceae Par

ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

48

hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

49

cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

50

ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 43: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

43

Balanitaceae Balanopaceae Balanophoraceae Balsameaceae Balsaminaceae Bambusa

ceae Barbaceniaceae BarbeuiaceaeBarbeyaceae Barclayaceae Barringtoniaceae Base

llaceae Bataceae Baueraceae Bauhiniaceae Baxteriaceae BegoniaceaeBehniaceae B

elangeraceae Belloniaceae Bembiciaceae Berberidaceae Berberidopsidaceae Berryac

eaee BersamaceaeBertyaceae Berzeliaceae Besleriaceae Betaceae Betulaceae Biebe

rsteiniaceae Bifariaceae Bignoniaceae BischofiaceaeBixaceae Blakeaceae Blandfordia

ceae Blattiaceae Blepharocaryaceae Blitaceae Blyxaceae Boerlagellaceae Bolivaracea

eBombacaceae Bonnetiaceae Bontiaceae Boopidaceae Boraginaceae Borassaceae B

oroniaceae BoryaceaeBotryodendraceae Bougainvilleaceae Boweniaceae Brassicacea

e Brachycaulaceae Bretschneideraceae BrexiaceaeBromeliaceae Brownlowiaceae Br

unelliaceae Bruniaceae Brunoniaceae Brunsvigiaceae Bryoniaceae BuchneraceaeBuci

daceae Buddlejaceae Buglossaceae Bulbocodiaceae Bumeliaceae Bupleuraceae Burc

hardiaceae BurmanniaceaeBurseraceae Butneriaceae Butomaceae Buxaceae Byblida

ceae Byttneriaceae

Cabombaceae Cacaoaceae Cactaceae Caesalpiniaceae Caladiaceae Calamaceae Calce

olariaceae CalectasiaceaeCalendulaceae Callaceae Callicomaceae Calligonaceae Calli

traceae Callitrichaceae Calochortaceae CalophyllaceaeCalthaceae Calycanthaceae Ca

lyceraceae Cambogiaceae Camelliaceae Campanulaceae CampynemataceaeCanacom

yricaceae Canellaceae Cannabaceae Cannaceae Canopodaceae Canotiaceae Cansjera

ceae CapparaceaeCaprariaceae Caprifoliaceae Cardiopteridaceae Carduaceae Carica

ceae Carissaceae Carlemanniaceae CarpinaceaeCarpodetaceae Cartonemataceae Ca

ryocaraceae Caryophyllaceae Casearieaceae Cassiaceae CassipoureaceaeCassythacea

e Castaneaceae Castelaceae Casuarinaceae Catesbaeaceae Cathedraceae Cecropiace

ae CedraceaeCedrelaceae Celastraceae Celosiaceae Celtidaceae Centaureaceae Cen

trolepidaceae Centroplacaceae CepaceaeCephalanthaceae Cephalotaceae Cephalota

xaceae Cerastiaceae Ceratoniaceae Ceratophyllaceae CerberaceaeCercidiphyllaceae

Cercocarpaceae Cercodiaceae Cereaceae Cerinthaceae Ceroxylaceae Cestraceae Cev

alliaceaeChailletiaceae Chamaedoraceae Chamaemoraceae Chamelauciaceae Chaun

ochitonaceae Chelidoniaceae ChelonaceaeChenopodiaceae Chimonanthaceae Chingi

thamnaceae Chionographidaceae Chiranthodendraceae ChironiaceaeChlaenaceae Ch

loanthaceae Chloranthaceae Chloridaceae Chlorogalaceae Chrysobalanaceae Cicerac

eae CichoriaceaeCimicifugaceae Cinchoniaceae Circaeaceae Circaeasteraceae Cissac

eae Cistaceae Citraceae Clematidaceae CleomaceaeClethraceae Clusiaceae Cneorac

eae Cnestidaceae Cnicaceae Cobaeaceae Cochlospermaceae Cocosaceae Coffeaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

44

Colchicaceae Coleogynaceae Columelliaceae Combretaceae Commelinaceae Comocl

adiaceae Compositae CompsoaceaeConantheraceae Connaraceae Conostylidaceae C

onvallariaceae Convolvulaceae Cordiaceae CoreopsidaceaeCoriandraceae Coriariacea

e Coridaceae Corispermaceae Cornaceae Corokiaceae Coronillaceae Correaceae Cor

rigiolaceaeCorsiaceae Corydalaceae Corylaceae Corynocarpaceae Coryphaceae Cost

aceae Cotyledonaceae CoulaceaeCoutariaceae Coutoubiaceae Crassulaceae Crescen

tiaceae Cressaceae Crinaceae Crocaceae CroomiaceaeCrossosomataceae Crotonacea

e Cruciferae Cryptaceae Crypteroniaceae Cryptocorynaceae CryptomeriaceaeCtenol

ophonaceae Cucurbitaceae Cunninghamiaceae Cunoniaceae Cupressaceae Curcumac

eae Curtisiaceae CuscutaceaeCyananthaceae Cyanastraceae Cynanchaceae Cyanella

ceae Cycadaceae Cyclanthaceae CyclantheraceaeCyclocheilaceae Cydoniaceae Cymo

doceaceae Cynaraceae Cynocrambaceae Cynomoriaceae Cyperaceae CyphiaceaeCyp

hocarpaceae Cypripediaceae Cyrillaceae Cyrtandraceae Cyrtanthaceae Cytinaceae

Dacrycarpaceae Dactylanthaceae Damasoniaceae Daphnaceae Daphniphyllaceae Das

ypogonaceae DatiscaceaeDaturaceae Daucaceae Davidiaceae Davidsoniaceae Decais

neaceae Deeringiaceae Degeneriaceae DelphiniaceaeDendrophthoaceae Columelliac

eae Detariaceae Dialypetalanthaceae Dianellaceae Dianthaceae DiapensiaceaeDicha

petalaceae Dichondraceae Diclidantheraceae Dicrastylidiaceae Dictamnaceae Didiere

aceae DidymelaceaeDidymocarpaceae Diegodendraceae Diervillaceae Digitalidaceae

e Dilatridaceae Dilleniaceae Dioaceae DionaeaceaeDioncophyllaceae Dioscoreaceae

Diosmaceae Diospyraceae Dipentodontaceae Diphylleiaceae Diplolaenaceae Dipsaca

ceaeDipterocarpaceae Dirachmaceae Disanthaceae Diselmaceae Dodonaeaceae Do

mbeyaceae Donatiaceae DorsteniaceaeDortmannaceae Doryanthaceae Drabaceae D

racaenaceae Dracontiaceae Drimyidaceae Droseraceae DrosophyllaceaeDryadaceae

Duabangaceae Duckeodendraceae Dulongiaceae Durantaceae Durionaceae Dysphani

aceae

Ebenaceae Eccremidaceae Ecdeiocoleaceae Echiaceae Echinopaceae Ehretiaceae Ela

eagnaceae ElaeocarpaceaeElegiaceae Elatinaceae Ellisiaceae Ellisiophyllaceae Elodea

ceae Elytranthaceae Embeliaceae Emblingiaceae EmmotaceaeEmpetraceae Encepha

lartaceae Engelhardtiaceae Enhalaceae Epacridaceae Ephedraceae Ephemeranthacea

eEpilobiaceae Epimediaceae Eragrostidaceae Eremolepidaceae Eremosynaceae Erica

ceae Erinaceae EriocaulaceaeEriogonaceae Eriospermaceae Erodiaceae Erycibaceae

Eryngiaceae Erysimaceae Erythroniaceae ErythropalaceaeErythrospermaceae Erythro

xylaceae Escalloniaceae Eschscholziaceae Eucomidaceae Eucommiaceae Eucryphiace

aeEuonymaceae Eupatoriaceae Euphorbiaceae Euphrasiaceae Euphroniaceae Eupo

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

45

matiaceae Eupteleaceae EuryalaceaeEustrephaceae Euthemidaceae Eutocaceae Exb

ucklandiaceae Exocarpaceae

Fabaceae Fagaceae Falcatifoliaceae Ferulaceae Festucaceae Ficaceae Ficoidaceae Fit

zroyaceae FlacourtiaceaeFlagellariaceae Flindersiaceae Foetidiaceae Forestieraceae

Fothergillaceae Fouquieriaceae Fragariaceae FrancoaceaeFrangulaceae Frankeniacea

e Fraxinaceae Fraxinellaceae Freycinetiaceae Fritillariaceae Fuchsiaceae Fumariacea

eFunkiaceae

Gaiadendraceae Galacaceae Galanthaceae Galaxiaceae Galedupaceae Galeniaceae G

aliaceae Garciniaceae GardeniaceaeGardneriaceae Garryaceae Geissolomataceae Ge

itonoplesiaceae Gelsemiaceae Geniostomaceae GentianaceaeGeonomataceae Geosir

idaceae Geraniaceae Gerrardinaceae Gesneriaceae Gethyllidaceae Gilliesiaceae Gina

lloaceaeGinkgoaceae Gisekiaceae Gladiolaceae Glaucidiaceae Glechomaceae Glinace

ae Globulariaceae Gnaphaliaceae GnetaceaeGoetzeaceae Gomortegaceae Gomphiac

eae Gomphrenaceae Gonystylaceae Goodeniaceae Gordoniaceae GouaniaceaeGoup

iaceae Gramineae Gratiolaceae Grewiaceae Greyiaceae Grielaceae Griseliniaceae Gr

onoviaceae GrossulariaceaeGrubbiaceae Guaiacanaceae Guamatelaceae Guettardac

eae Gunneraceae Gustaviaceae Guttiferae GyrocarpaceaeGyrostemonaceae

Hachetteaceae Haemanthaceae Haemodoraceae Halesiaceae Hallieraceae Halocarpa

ceae Halophilaceae HalophytaceaeHaloragaceae Hamamelidaceae Hanguanaceae H

aptanthaceae Harmandaceae Hebenstretiaceae HectorellaceaeHederaceae Hedyosm

aceae Hedyotidaceae Hedysaraceae Heisteriaceae Heleniaceae Heliamphoraceae H

elianthaceaeHelianthemaceae Helichrysaceae Heliconiaceae Helicteraceae Heliotropi

aceae Helleboraceae Heloniadaceae HelosidaceaeHelwingiaceae Hemerocallidaceae

Hemimeridaceae Hemiphylacaceae Henriqueziaceae Henslowiaceae Hermanniaceae

Hernandiaceae Herniariaceae Herreriaceae Hesperocallidaceae Heterantheraceae He

teropyxidaceae HeterostylaceaeHibbertiaceae Hibiscaceae Hilleriaceae Himantandra

ceae Hippocastanaceae Hippocrateaceae HippomanaceaeHippophaeaceae Hippurida

ceae Hirtellaceae Holacanthaceae Homaliaceae Hopkinsiaceae Hoplestigmataceae H

ordeaceaeHornschuchiaceae Hortensiaceae Hortoniaceae Hostaceae Houstoniaceae

Huaceae Huerteaceae HugoniaceaeHumbertiaceae Humiriaceae Hyacinthaceae Hyd

atellaceae Hydnoraceae Hydrangeaceae Hydrastidaceae HydrillaceaeHydrocharitacea

e Hydrocotylaceae Hydrogetonaceae Hydroleaceae Hydropeltidaceae Hydrophylacac

eae HydrophyllaceaeHydrostachyaceae Hymenocardiaceae Hypecoaceae Hyperanth

eraceae Hypericaceae Hypopityaceae HypoxidaceaeHypseocharitaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

46

Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta

ceae ImperatoriaceaeInocarpaceae Inulaceae Ionidiaceae Iriarteaceae Iridaceae Irvi

ngiaceae Isnardiaceae Isophysidaceae IteaceaeStrasburgeriaceae Ixiaceae Ixioliriace

ae Ixonanthaceae

Jalapaceae Jamboliferaceae Japonoliriaceae Jasionaceae Jasminiaceae Johnsoniaceae

Joinvilleaceae JuglandaceaeJulianiaceae Juncaceae Juncaginaceae Juniperaceae Jus

siaeaceae Justiciaceae

Kadsuraceae Kaliphoraceae Kalmiaceae Kaniaceae Kiggelariaceae Kingdoniaceae Kin

giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae

Krameriaceae

Labiatae Lacandoniaceae Lachenaliaceae Lacistemataceae Lactoridaceae Lactucaceae

Lagerstroemiaceaee LamiaceaeLampsanaceae Lanariaceae Langsdorffiaceae Lantan

aceae Lapageriaceae Lardizabalaceae Lasiopetalaceae LauraceaeLawsoniaceae Laxm

anniaceae Lecythidaceae Ledaceae Ledocarpaceae Leeaceae Leguminosae Leitneriac

eae LemnaceaeLennoaceae Lentibulariaceae Lentiscaceae Leoniaceae Leonticaceae

Lepidobotryaceae LepidocarpaceaeLepidocaryaceae Lepidocerataceae Leptaulaceae

Leptospermaceae Lepturaceae Lepuropetalaceae LeuctenbergiaceaeLibocedraceae

Licaniaceae Ligustraceae Lilaeaceae Liliaceae Limeaceae Limnanthaceae Limnocharit

aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace

ae Linderniaceae Lindleyaceae Linnaeaceae LippayaceaeLiriaceae Liriodendraceae Li

ssocarpaceae Littorellaceae Loasaceae Lobeliaceaee Loganiaceae LomandraceaeLoni

ceraceae Lopeziaceae Lophiocarpaceae Lophiolaceae Lophiraceae Lophophytaceae L

ophopyxidaceae LoranthaceaeLotaceae Lowiaceae Lupulaceaee Luxemburgiaceae Lu

zuriagaceae Lychnidaceae Lyciaceae LyginiaceaeLygodisodeaceae Lysimachiaceae Ly

thraceae

Macarisiaceae Mackinlayaceae Madiaceae Maesaceae Magnoliaceae Malaceae Male

sherbiaceae MalortieaceaeMalpighiaceae Malvaceae Mangiaceae Manicariaceae Ma

paniaceae Marantaceae Marathraceae MarcgraviaceaeMartyniaceae Mastixiaceae

Matricariaceae Maundiaceae Mayacaceae Medeolaceae Medusagynaceae Peridiscac

eaeMelaleucaceae Melampyraceae Melanophyllaceae Melanthiaceae Melastomatac

eae Meliaceae Melianthaceae MelicaceaeMeliosmaceae Melittidaceae Melochiacea

e Memecylaceae Mendonciaceae Menispermaceae Menthaceae MenyanthaceaeMe

nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

47

Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

Nypaceae Nyssaceae

Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

ceae Papilionaceae Pappophoraceae PapyraceaeParacryphiaceae Parasitaxaceae Par

ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

48

hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

49

cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

50

ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
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PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

44

Colchicaceae Coleogynaceae Columelliaceae Combretaceae Commelinaceae Comocl

adiaceae Compositae CompsoaceaeConantheraceae Connaraceae Conostylidaceae C

onvallariaceae Convolvulaceae Cordiaceae CoreopsidaceaeCoriandraceae Coriariacea

e Coridaceae Corispermaceae Cornaceae Corokiaceae Coronillaceae Correaceae Cor

rigiolaceaeCorsiaceae Corydalaceae Corylaceae Corynocarpaceae Coryphaceae Cost

aceae Cotyledonaceae CoulaceaeCoutariaceae Coutoubiaceae Crassulaceae Crescen

tiaceae Cressaceae Crinaceae Crocaceae CroomiaceaeCrossosomataceae Crotonacea

e Cruciferae Cryptaceae Crypteroniaceae Cryptocorynaceae CryptomeriaceaeCtenol

ophonaceae Cucurbitaceae Cunninghamiaceae Cunoniaceae Cupressaceae Curcumac

eae Curtisiaceae CuscutaceaeCyananthaceae Cyanastraceae Cynanchaceae Cyanella

ceae Cycadaceae Cyclanthaceae CyclantheraceaeCyclocheilaceae Cydoniaceae Cymo

doceaceae Cynaraceae Cynocrambaceae Cynomoriaceae Cyperaceae CyphiaceaeCyp

hocarpaceae Cypripediaceae Cyrillaceae Cyrtandraceae Cyrtanthaceae Cytinaceae

Dacrycarpaceae Dactylanthaceae Damasoniaceae Daphnaceae Daphniphyllaceae Das

ypogonaceae DatiscaceaeDaturaceae Daucaceae Davidiaceae Davidsoniaceae Decais

neaceae Deeringiaceae Degeneriaceae DelphiniaceaeDendrophthoaceae Columelliac

eae Detariaceae Dialypetalanthaceae Dianellaceae Dianthaceae DiapensiaceaeDicha

petalaceae Dichondraceae Diclidantheraceae Dicrastylidiaceae Dictamnaceae Didiere

aceae DidymelaceaeDidymocarpaceae Diegodendraceae Diervillaceae Digitalidaceae

e Dilatridaceae Dilleniaceae Dioaceae DionaeaceaeDioncophyllaceae Dioscoreaceae

Diosmaceae Diospyraceae Dipentodontaceae Diphylleiaceae Diplolaenaceae Dipsaca

ceaeDipterocarpaceae Dirachmaceae Disanthaceae Diselmaceae Dodonaeaceae Do

mbeyaceae Donatiaceae DorsteniaceaeDortmannaceae Doryanthaceae Drabaceae D

racaenaceae Dracontiaceae Drimyidaceae Droseraceae DrosophyllaceaeDryadaceae

Duabangaceae Duckeodendraceae Dulongiaceae Durantaceae Durionaceae Dysphani

aceae

Ebenaceae Eccremidaceae Ecdeiocoleaceae Echiaceae Echinopaceae Ehretiaceae Ela

eagnaceae ElaeocarpaceaeElegiaceae Elatinaceae Ellisiaceae Ellisiophyllaceae Elodea

ceae Elytranthaceae Embeliaceae Emblingiaceae EmmotaceaeEmpetraceae Encepha

lartaceae Engelhardtiaceae Enhalaceae Epacridaceae Ephedraceae Ephemeranthacea

eEpilobiaceae Epimediaceae Eragrostidaceae Eremolepidaceae Eremosynaceae Erica

ceae Erinaceae EriocaulaceaeEriogonaceae Eriospermaceae Erodiaceae Erycibaceae

Eryngiaceae Erysimaceae Erythroniaceae ErythropalaceaeErythrospermaceae Erythro

xylaceae Escalloniaceae Eschscholziaceae Eucomidaceae Eucommiaceae Eucryphiace

aeEuonymaceae Eupatoriaceae Euphorbiaceae Euphrasiaceae Euphroniaceae Eupo

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

45

matiaceae Eupteleaceae EuryalaceaeEustrephaceae Euthemidaceae Eutocaceae Exb

ucklandiaceae Exocarpaceae

Fabaceae Fagaceae Falcatifoliaceae Ferulaceae Festucaceae Ficaceae Ficoidaceae Fit

zroyaceae FlacourtiaceaeFlagellariaceae Flindersiaceae Foetidiaceae Forestieraceae

Fothergillaceae Fouquieriaceae Fragariaceae FrancoaceaeFrangulaceae Frankeniacea

e Fraxinaceae Fraxinellaceae Freycinetiaceae Fritillariaceae Fuchsiaceae Fumariacea

eFunkiaceae

Gaiadendraceae Galacaceae Galanthaceae Galaxiaceae Galedupaceae Galeniaceae G

aliaceae Garciniaceae GardeniaceaeGardneriaceae Garryaceae Geissolomataceae Ge

itonoplesiaceae Gelsemiaceae Geniostomaceae GentianaceaeGeonomataceae Geosir

idaceae Geraniaceae Gerrardinaceae Gesneriaceae Gethyllidaceae Gilliesiaceae Gina

lloaceaeGinkgoaceae Gisekiaceae Gladiolaceae Glaucidiaceae Glechomaceae Glinace

ae Globulariaceae Gnaphaliaceae GnetaceaeGoetzeaceae Gomortegaceae Gomphiac

eae Gomphrenaceae Gonystylaceae Goodeniaceae Gordoniaceae GouaniaceaeGoup

iaceae Gramineae Gratiolaceae Grewiaceae Greyiaceae Grielaceae Griseliniaceae Gr

onoviaceae GrossulariaceaeGrubbiaceae Guaiacanaceae Guamatelaceae Guettardac

eae Gunneraceae Gustaviaceae Guttiferae GyrocarpaceaeGyrostemonaceae

Hachetteaceae Haemanthaceae Haemodoraceae Halesiaceae Hallieraceae Halocarpa

ceae Halophilaceae HalophytaceaeHaloragaceae Hamamelidaceae Hanguanaceae H

aptanthaceae Harmandaceae Hebenstretiaceae HectorellaceaeHederaceae Hedyosm

aceae Hedyotidaceae Hedysaraceae Heisteriaceae Heleniaceae Heliamphoraceae H

elianthaceaeHelianthemaceae Helichrysaceae Heliconiaceae Helicteraceae Heliotropi

aceae Helleboraceae Heloniadaceae HelosidaceaeHelwingiaceae Hemerocallidaceae

Hemimeridaceae Hemiphylacaceae Henriqueziaceae Henslowiaceae Hermanniaceae

Hernandiaceae Herniariaceae Herreriaceae Hesperocallidaceae Heterantheraceae He

teropyxidaceae HeterostylaceaeHibbertiaceae Hibiscaceae Hilleriaceae Himantandra

ceae Hippocastanaceae Hippocrateaceae HippomanaceaeHippophaeaceae Hippurida

ceae Hirtellaceae Holacanthaceae Homaliaceae Hopkinsiaceae Hoplestigmataceae H

ordeaceaeHornschuchiaceae Hortensiaceae Hortoniaceae Hostaceae Houstoniaceae

Huaceae Huerteaceae HugoniaceaeHumbertiaceae Humiriaceae Hyacinthaceae Hyd

atellaceae Hydnoraceae Hydrangeaceae Hydrastidaceae HydrillaceaeHydrocharitacea

e Hydrocotylaceae Hydrogetonaceae Hydroleaceae Hydropeltidaceae Hydrophylacac

eae HydrophyllaceaeHydrostachyaceae Hymenocardiaceae Hypecoaceae Hyperanth

eraceae Hypericaceae Hypopityaceae HypoxidaceaeHypseocharitaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

46

Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta

ceae ImperatoriaceaeInocarpaceae Inulaceae Ionidiaceae Iriarteaceae Iridaceae Irvi

ngiaceae Isnardiaceae Isophysidaceae IteaceaeStrasburgeriaceae Ixiaceae Ixioliriace

ae Ixonanthaceae

Jalapaceae Jamboliferaceae Japonoliriaceae Jasionaceae Jasminiaceae Johnsoniaceae

Joinvilleaceae JuglandaceaeJulianiaceae Juncaceae Juncaginaceae Juniperaceae Jus

siaeaceae Justiciaceae

Kadsuraceae Kaliphoraceae Kalmiaceae Kaniaceae Kiggelariaceae Kingdoniaceae Kin

giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae

Krameriaceae

Labiatae Lacandoniaceae Lachenaliaceae Lacistemataceae Lactoridaceae Lactucaceae

Lagerstroemiaceaee LamiaceaeLampsanaceae Lanariaceae Langsdorffiaceae Lantan

aceae Lapageriaceae Lardizabalaceae Lasiopetalaceae LauraceaeLawsoniaceae Laxm

anniaceae Lecythidaceae Ledaceae Ledocarpaceae Leeaceae Leguminosae Leitneriac

eae LemnaceaeLennoaceae Lentibulariaceae Lentiscaceae Leoniaceae Leonticaceae

Lepidobotryaceae LepidocarpaceaeLepidocaryaceae Lepidocerataceae Leptaulaceae

Leptospermaceae Lepturaceae Lepuropetalaceae LeuctenbergiaceaeLibocedraceae

Licaniaceae Ligustraceae Lilaeaceae Liliaceae Limeaceae Limnanthaceae Limnocharit

aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace

ae Linderniaceae Lindleyaceae Linnaeaceae LippayaceaeLiriaceae Liriodendraceae Li

ssocarpaceae Littorellaceae Loasaceae Lobeliaceaee Loganiaceae LomandraceaeLoni

ceraceae Lopeziaceae Lophiocarpaceae Lophiolaceae Lophiraceae Lophophytaceae L

ophopyxidaceae LoranthaceaeLotaceae Lowiaceae Lupulaceaee Luxemburgiaceae Lu

zuriagaceae Lychnidaceae Lyciaceae LyginiaceaeLygodisodeaceae Lysimachiaceae Ly

thraceae

Macarisiaceae Mackinlayaceae Madiaceae Maesaceae Magnoliaceae Malaceae Male

sherbiaceae MalortieaceaeMalpighiaceae Malvaceae Mangiaceae Manicariaceae Ma

paniaceae Marantaceae Marathraceae MarcgraviaceaeMartyniaceae Mastixiaceae

Matricariaceae Maundiaceae Mayacaceae Medeolaceae Medusagynaceae Peridiscac

eaeMelaleucaceae Melampyraceae Melanophyllaceae Melanthiaceae Melastomatac

eae Meliaceae Melianthaceae MelicaceaeMeliosmaceae Melittidaceae Melochiacea

e Memecylaceae Mendonciaceae Menispermaceae Menthaceae MenyanthaceaeMe

nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

47

Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

Nypaceae Nyssaceae

Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

ceae Papilionaceae Pappophoraceae PapyraceaeParacryphiaceae Parasitaxaceae Par

ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

48

hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

49

cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

50

ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
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PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

45

matiaceae Eupteleaceae EuryalaceaeEustrephaceae Euthemidaceae Eutocaceae Exb

ucklandiaceae Exocarpaceae

Fabaceae Fagaceae Falcatifoliaceae Ferulaceae Festucaceae Ficaceae Ficoidaceae Fit

zroyaceae FlacourtiaceaeFlagellariaceae Flindersiaceae Foetidiaceae Forestieraceae

Fothergillaceae Fouquieriaceae Fragariaceae FrancoaceaeFrangulaceae Frankeniacea

e Fraxinaceae Fraxinellaceae Freycinetiaceae Fritillariaceae Fuchsiaceae Fumariacea

eFunkiaceae

Gaiadendraceae Galacaceae Galanthaceae Galaxiaceae Galedupaceae Galeniaceae G

aliaceae Garciniaceae GardeniaceaeGardneriaceae Garryaceae Geissolomataceae Ge

itonoplesiaceae Gelsemiaceae Geniostomaceae GentianaceaeGeonomataceae Geosir

idaceae Geraniaceae Gerrardinaceae Gesneriaceae Gethyllidaceae Gilliesiaceae Gina

lloaceaeGinkgoaceae Gisekiaceae Gladiolaceae Glaucidiaceae Glechomaceae Glinace

ae Globulariaceae Gnaphaliaceae GnetaceaeGoetzeaceae Gomortegaceae Gomphiac

eae Gomphrenaceae Gonystylaceae Goodeniaceae Gordoniaceae GouaniaceaeGoup

iaceae Gramineae Gratiolaceae Grewiaceae Greyiaceae Grielaceae Griseliniaceae Gr

onoviaceae GrossulariaceaeGrubbiaceae Guaiacanaceae Guamatelaceae Guettardac

eae Gunneraceae Gustaviaceae Guttiferae GyrocarpaceaeGyrostemonaceae

Hachetteaceae Haemanthaceae Haemodoraceae Halesiaceae Hallieraceae Halocarpa

ceae Halophilaceae HalophytaceaeHaloragaceae Hamamelidaceae Hanguanaceae H

aptanthaceae Harmandaceae Hebenstretiaceae HectorellaceaeHederaceae Hedyosm

aceae Hedyotidaceae Hedysaraceae Heisteriaceae Heleniaceae Heliamphoraceae H

elianthaceaeHelianthemaceae Helichrysaceae Heliconiaceae Helicteraceae Heliotropi

aceae Helleboraceae Heloniadaceae HelosidaceaeHelwingiaceae Hemerocallidaceae

Hemimeridaceae Hemiphylacaceae Henriqueziaceae Henslowiaceae Hermanniaceae

Hernandiaceae Herniariaceae Herreriaceae Hesperocallidaceae Heterantheraceae He

teropyxidaceae HeterostylaceaeHibbertiaceae Hibiscaceae Hilleriaceae Himantandra

ceae Hippocastanaceae Hippocrateaceae HippomanaceaeHippophaeaceae Hippurida

ceae Hirtellaceae Holacanthaceae Homaliaceae Hopkinsiaceae Hoplestigmataceae H

ordeaceaeHornschuchiaceae Hortensiaceae Hortoniaceae Hostaceae Houstoniaceae

Huaceae Huerteaceae HugoniaceaeHumbertiaceae Humiriaceae Hyacinthaceae Hyd

atellaceae Hydnoraceae Hydrangeaceae Hydrastidaceae HydrillaceaeHydrocharitacea

e Hydrocotylaceae Hydrogetonaceae Hydroleaceae Hydropeltidaceae Hydrophylacac

eae HydrophyllaceaeHydrostachyaceae Hymenocardiaceae Hypecoaceae Hyperanth

eraceae Hypericaceae Hypopityaceae HypoxidaceaeHypseocharitaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

46

Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta

ceae ImperatoriaceaeInocarpaceae Inulaceae Ionidiaceae Iriarteaceae Iridaceae Irvi

ngiaceae Isnardiaceae Isophysidaceae IteaceaeStrasburgeriaceae Ixiaceae Ixioliriace

ae Ixonanthaceae

Jalapaceae Jamboliferaceae Japonoliriaceae Jasionaceae Jasminiaceae Johnsoniaceae

Joinvilleaceae JuglandaceaeJulianiaceae Juncaceae Juncaginaceae Juniperaceae Jus

siaeaceae Justiciaceae

Kadsuraceae Kaliphoraceae Kalmiaceae Kaniaceae Kiggelariaceae Kingdoniaceae Kin

giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae

Krameriaceae

Labiatae Lacandoniaceae Lachenaliaceae Lacistemataceae Lactoridaceae Lactucaceae

Lagerstroemiaceaee LamiaceaeLampsanaceae Lanariaceae Langsdorffiaceae Lantan

aceae Lapageriaceae Lardizabalaceae Lasiopetalaceae LauraceaeLawsoniaceae Laxm

anniaceae Lecythidaceae Ledaceae Ledocarpaceae Leeaceae Leguminosae Leitneriac

eae LemnaceaeLennoaceae Lentibulariaceae Lentiscaceae Leoniaceae Leonticaceae

Lepidobotryaceae LepidocarpaceaeLepidocaryaceae Lepidocerataceae Leptaulaceae

Leptospermaceae Lepturaceae Lepuropetalaceae LeuctenbergiaceaeLibocedraceae

Licaniaceae Ligustraceae Lilaeaceae Liliaceae Limeaceae Limnanthaceae Limnocharit

aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace

ae Linderniaceae Lindleyaceae Linnaeaceae LippayaceaeLiriaceae Liriodendraceae Li

ssocarpaceae Littorellaceae Loasaceae Lobeliaceaee Loganiaceae LomandraceaeLoni

ceraceae Lopeziaceae Lophiocarpaceae Lophiolaceae Lophiraceae Lophophytaceae L

ophopyxidaceae LoranthaceaeLotaceae Lowiaceae Lupulaceaee Luxemburgiaceae Lu

zuriagaceae Lychnidaceae Lyciaceae LyginiaceaeLygodisodeaceae Lysimachiaceae Ly

thraceae

Macarisiaceae Mackinlayaceae Madiaceae Maesaceae Magnoliaceae Malaceae Male

sherbiaceae MalortieaceaeMalpighiaceae Malvaceae Mangiaceae Manicariaceae Ma

paniaceae Marantaceae Marathraceae MarcgraviaceaeMartyniaceae Mastixiaceae

Matricariaceae Maundiaceae Mayacaceae Medeolaceae Medusagynaceae Peridiscac

eaeMelaleucaceae Melampyraceae Melanophyllaceae Melanthiaceae Melastomatac

eae Meliaceae Melianthaceae MelicaceaeMeliosmaceae Melittidaceae Melochiacea

e Memecylaceae Mendonciaceae Menispermaceae Menthaceae MenyanthaceaeMe

nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

47

Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

Nypaceae Nyssaceae

Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

ceae Papilionaceae Pappophoraceae PapyraceaeParacryphiaceae Parasitaxaceae Par

ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

48

hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

49

cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

50

ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 46: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

46

Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta

ceae ImperatoriaceaeInocarpaceae Inulaceae Ionidiaceae Iriarteaceae Iridaceae Irvi

ngiaceae Isnardiaceae Isophysidaceae IteaceaeStrasburgeriaceae Ixiaceae Ixioliriace

ae Ixonanthaceae

Jalapaceae Jamboliferaceae Japonoliriaceae Jasionaceae Jasminiaceae Johnsoniaceae

Joinvilleaceae JuglandaceaeJulianiaceae Juncaceae Juncaginaceae Juniperaceae Jus

siaeaceae Justiciaceae

Kadsuraceae Kaliphoraceae Kalmiaceae Kaniaceae Kiggelariaceae Kingdoniaceae Kin

giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae

Krameriaceae

Labiatae Lacandoniaceae Lachenaliaceae Lacistemataceae Lactoridaceae Lactucaceae

Lagerstroemiaceaee LamiaceaeLampsanaceae Lanariaceae Langsdorffiaceae Lantan

aceae Lapageriaceae Lardizabalaceae Lasiopetalaceae LauraceaeLawsoniaceae Laxm

anniaceae Lecythidaceae Ledaceae Ledocarpaceae Leeaceae Leguminosae Leitneriac

eae LemnaceaeLennoaceae Lentibulariaceae Lentiscaceae Leoniaceae Leonticaceae

Lepidobotryaceae LepidocarpaceaeLepidocaryaceae Lepidocerataceae Leptaulaceae

Leptospermaceae Lepturaceae Lepuropetalaceae LeuctenbergiaceaeLibocedraceae

Licaniaceae Ligustraceae Lilaeaceae Liliaceae Limeaceae Limnanthaceae Limnocharit

aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace

ae Linderniaceae Lindleyaceae Linnaeaceae LippayaceaeLiriaceae Liriodendraceae Li

ssocarpaceae Littorellaceae Loasaceae Lobeliaceaee Loganiaceae LomandraceaeLoni

ceraceae Lopeziaceae Lophiocarpaceae Lophiolaceae Lophiraceae Lophophytaceae L

ophopyxidaceae LoranthaceaeLotaceae Lowiaceae Lupulaceaee Luxemburgiaceae Lu

zuriagaceae Lychnidaceae Lyciaceae LyginiaceaeLygodisodeaceae Lysimachiaceae Ly

thraceae

Macarisiaceae Mackinlayaceae Madiaceae Maesaceae Magnoliaceae Malaceae Male

sherbiaceae MalortieaceaeMalpighiaceae Malvaceae Mangiaceae Manicariaceae Ma

paniaceae Marantaceae Marathraceae MarcgraviaceaeMartyniaceae Mastixiaceae

Matricariaceae Maundiaceae Mayacaceae Medeolaceae Medusagynaceae Peridiscac

eaeMelaleucaceae Melampyraceae Melanophyllaceae Melanthiaceae Melastomatac

eae Meliaceae Melianthaceae MelicaceaeMeliosmaceae Melittidaceae Melochiacea

e Memecylaceae Mendonciaceae Menispermaceae Menthaceae MenyanthaceaeMe

nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

47

Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

Nypaceae Nyssaceae

Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

ceae Papilionaceae Pappophoraceae PapyraceaeParacryphiaceae Parasitaxaceae Par

ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

48

hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

49

cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

50

ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 47: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

47

Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth

eaceae Microbiotaceae Microcachrydaceae MicrocycadaceaeMiliaceae Milulaceae

Mimosaceae Mirabilidaceaee Misodendraceae Mitrastemonaceae Miyoshiaceae Mo

deccaceaeMolluginaceae Monimiaceae Monodoraceae Monotaceae Monotropaceae

Montiaceae Montiniaceae Moraceae MorinaceaeMoringaceae Mouririaceae Mout

abeaceae Muntingiaceae Musaceae Mutisiaceae Myodocarpaceae MyoporaceaeMyr

icaceae Myriophyllaceae Myristicaceae Myrobalanaceae Myrothamnaceae Myrrhinia

ceae Myrsinaceae MyrtaceaeMystropetalaceae

Nageiaceae Najadaceae Nandinaceae Napoleonaceae Nartheciaceae Narcissiaceaee

Nardaceae NassauviaceaeNaucleaceae Nectaropetalaceae Neilliaceae Nelsoniaceae

Nelumbonaceae Nemacladaceae NeocallitropsidaceaeNeottiaceae Nepenthaceae Ne

petaceae Nesogenaceae Neuradaceae Neuwiediaceae Nhandirobaceae Nicotaniacea

eNigellaceae Nitrariaceae Nolanaceae Nolinaceae Nonateliaceae Nopaleaceae Noth

ofagaceae Nupharaceae NuytsiaceaeNyctaginaceae Nyctanthaceae Nymphaeaceae

Nypaceae Nyssaceae

Obolariaceae Ochnaceae Ochranthaceae Octoknemaceae Oenotheraceae Oftiaceae

Olacaceae Oleaceae OliniaceaeOnagraceae Oncothecaceae Onosmaceae Opercula

riaceae Ophiopogonaceae Ophioxylaceae Ophiraceae OpiliaceaeOporanthaceae Opu

ntiaceae Orchidaceae Ornithogalaceae Ornithrophaceae Orobanchaceae Orontiaceae

OrtegaceaeOryzaceae Osyridaceae Oxalidaceae Oxycladaceae Oxycoccaceae Oxysty

lidaceae

Pachysandraceae Pacouriaceae Paeoniaceae Pagamaeaceae Paivaeusaceae Palmae

Pancratiaceae PandaceaePandanaceae Pangiaceae Panicaceae Papaveraceae Papaya

ceae Papilionaceae Pappophoraceae PapyraceaeParacryphiaceae Parasitaxaceae Par

ianaceae Paridaceae Parnassiaceae Paronychiaceae Paropsiaceae ParrotiaceaeParth

eniaceae Passifloraceae Pastinacaceae Paulliniaceae Paulowniaceae Paviaceae Pecti

antiaceae PedaliaceaePedicularidaceae Peganaceae Peliosanthaceae Pellicieraceae P

enaeaceae Pennantiaceae PentadiplandraceaePentapetaceae Pentaphragmataceae P

entaphylacaceae Pentastemonaceae Penthoraceae Peperomiaceae PeraceaePerdicia

ceae Peridiscaceae Periplocaceae Peripterygiaceae Perseaceae Persicariaceae Peter

manniaceae PetiveriaceaePetreaceae Petrosaviaceae Phalaridaceae Phaleriaceae Ph

araceae Pharnaceaceae Phaseolaceae PhellinaceaePhelypaeaceae Pherosphaeraceae

Philadelphaceae Philesiaceae Philippodendraceae Philocrenaceae PhilydraceaePhoe

niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

48

hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

49

cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

50

ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 48: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

48

hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol

accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P

ilocarpaceae Pinaceae Pinguiculaceae Piperaceae Piriquetaceae Pisoniaceae Pistacia

ceae PistiaceaePittosporaceae Plagianthaceae Plagiopteraceae Plantaginaceae Plata

naceae Platycaryaceae PlatycladaceaePlatyspermataceae Plectroniaceae Pleurisanth

aceae Platystemonaceae Plocospermataceae Plumbaginaceae PlumeriaceaePoaceae

Podoaceae Podocarpaceae Podophyllaceae Podostemaceae Polemoniaceae Poliothy

rsidaceae PolpodaceaePolycarpaeaceae Polycnemaceae Polygalaceae Polygonaceae

Polygonanthaceae Polygonataceae PolyosmaceaePontederiaceae Poranaceae Porant

heraceae Portulacaceae Portulacariaceae Posidoniaceae PotaliaceaePotamogetonace

ae Potentillaceae Poteriaceae Pothaceae Pottingeriaceae Primulaceae Prioniaceae P

rionotaceaeProckiaceae Proteaceae Prumnopityaceae Prunaceae Pseliaceae Pseuda

nthaceae Pseudophoeniciaceae PsiloxylaceaePsittacanthaceae Psychotriaceae Psyllia

ceaee Ptaeroxylaceae Pteleaceae Pteleocarpaceae PteridophyllaceaePterisanthaceae

Pterostemonaceae Punicaceae Putranjivaceae Pyraceae Pyrolaceae

Quassiaceae Quercaceae Quiinaceae Quillajaceae Paracryphiaceae

Rafflesiaceae Ramondaceae Randiaceae Ranunculaceae Ranzaniaceae Rapateaceae

Raphanaceae ReaumuriaceaeResedaceae Restionaceae Retziaceae Rhabdodendracea

e Rhamnaceae Rhaptopetalaceae Rhexiaceae RhinanthaceaeRhipogonaceae Rhizoph

oraceae Rhodiolaceae Rhododendraceae Rhodolaenaceae Rhodoleiaceae Rhodorace

aeRhodotypaceae Rhoipteleaceae Rhopalocarpaceae Rhynchocalycaceae Rhynchoth

ecaceae Ribesiaceae RicinaceaeRicinocarpaceae Riviniaceae Roridulaceae Rosaceae

Rousseaceae Roxburghiaceae Rubiaceae Rumicaceae RuppiaceaeRuscaceae Rutacea

e

Sabalaceae Sabiaceae Sabiceaceae Saccharaceae Saccifoliaceae Sagoneaceae Salacia

ceae SalaxidaceaeSalazariaceae Salicaceae Salicorniaceae Salpiglosssidaceae Salsolac

eae Salvadoraceae Salviaceae SambucaceaeSamolaceae Samydaceae Sanguisorbace

ae Saniculaceae Sanseveriaceae Santalaceae Santolinaceae SapindaceaeSapotaceae

Sarcobataceae Sarcocaceae Sarcolaenaceae Sarcophytaceae Sarcospermataceae Sarc

ostigmataceaeSargentodoxaceae Sarraceniaceae Saurauiaceae Saururaceae Sauvage

siaceae Saxegotheaceae SaxifragaceaeScabiosaceae Scaevolaceae Scepaceae Scheuc

hzeriaceae Schinaceae Schisandraceae Schizolaenaceae SchlegeliaceaeSchoepfiaceae

Schreberaceae Sciadopityaceae Scillaceae Scirpaceae Scleranthaceae Sclerophylacac

eae ScoliopaceaeScopariaceae Scorodocarpaceae Scrophulariaceae Scutellariaceae S

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

49

cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

50

ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 49: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

49

cybaliaceae Scyphostegiaceae ScytopetalaceaeSebestenaceae Sedaceae Seguieriacea

e Selaginaceae Sempervivaceae Senecionaceae Sequoiaceae SerrulataceaeSesamace

ae Sesuviaceae Setchellanthaceae Sheadendraceae Sibthorpiaceae Silenaceae Simab

aceae SimaroubaceaeSimmondsiaceae Sinofranchetiaceae Siparunaceae Siphonanth

aceae Siphonodontaceae Sisymbriaceae SladeniaceaeSmeathmanniaceae Smilacacea

e Smyrniaceae Solanaceae Sonneratiaceae Soramiaceae Sorbaceae SoulameaceaeeS

parganiaceae Sparmanniaceae Spartinaceae Spatheliaceae Spergulaceae Sphaerosep

alaceae SphenocleaceaeSphenostemonaceae Spigeliaceae Spiniciaceae Spiraeaceae

Spiraeanthemaceae Spondiadaceae SporobolaceaeStachyuraceae Stackhousiaceae St

angeriaceae Stanleyaceae Stapeliaceae Staphyleaceae StaticaceaeStegnospermatace

ae Stellariaceae Stemonaceae Stemonuraceae Stenomeridaceae Sterculiaceae Stilagi

naceaeStilbaceae Stipaceae Stixaceae Strasburgeriaceae Stratiotaceae Strelitziaceae

Streptochaetaceae StrombosiaceaeStrumariaceae Strychnaceaee Stylidiaceae Stylob

asiaceae Stylocerataceae Stypheliaceae Styracaceae SurianaceaeSwartziaceae Swiet

eniaceae Symphoremataceae Symplocaceae Synechanthaceae Syringaceae

Taccaceae Taiwaniaceae Taktajaniaceae Talinaceae Tamaceae Tamaricaceae Tamari

ndaceae TamnaceaeTanacetaceae Tapisciaceae Taxaceae Taxodiaceae Tecophilaeac

eae Telephiaceae Tepuianthaceae TerminaliaceaeTernstroemiaceae Tetracarpaeacea

e Tetracentraceae Tetrachondraceae Tetraclinaceae Tetradiclidaceae Tetragoniaceae

Tetramelaceae Tetrameristaceae Tetrastylidiaceae Tetrathecaceae Thalassiaceae Th

alictraceae TheaceaeTheligonaceae Themidaceae Theobromataceae Theophrastacea

e Thesiaceae Thismiaceae Thlaspiaceae ThoaceaeThomandersiaceae Thujaceae Thuj

opsidaceae Thunbergiaceae Thurniaceae Thymelaeaceae Ticodendraceae TiliaceaeTi

llaeaceae Tillandsiaceae Tinaceae Tithymalaceae Tofieldiaceae Tormentillaceae Torr

eyaceae TorricelliaceaeTovariaceae Tradescantiaceae Euphorbiaceae Trapaceae Tra

pellaceae Tremandraceae Trewiaceae TribelaceaeTribulaceae Trichopodaceae Tricyr

tidaceae Triglochinaceaee Trigoniaceae Trilliaceae Trimeniaceae TriplobaceaeTriploc

hitonaceae Triplostegiaceae Tristichaceae Triticaceae Triuridaceae Trochodendracea

e TropaeolaceaeTulbaghiaceae Tulipaceae Tumboaceae Tupistraceae Turneraceae T

yphaceae

Uapacaceae Ullucaceae Ulmaceae Ulmariaceae Umbelliferae Unisemataceae Urticac

eae Utriculariaceae Uvulariaceae

Vacciniaceae Vahliaceae Valerianaceae Vallisneraceae Vanillaceae Velloziaceae Vera

traceae Verbascaceae VerbenaceaeVernicaceae Vernoniaceae Veronicaceae Viburna

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

50

ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 50: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

50

ceae Viciaceae Vincaceae Violaceae Viscaceae Vitaceae ViticaceaeVivianiaceae Voc

hysiaceae

Wachendorfiaceae Walleriaceae Wellingtoniaceae Wellstediaceae Welwitschiaceae

Widdringtoniaceae WillughbieaceaeWinteraceae Woffiaceae

Xanthiaceae Xanthophyllaceae Xanthorrhoeaceae Xeronemataceae Xerophyllaceae X

imeniaceae Xiphidiaceae Xyridaceae

Yuccaceae

Zamiaceae Zannichelliaceae Zanoniaceae Zanthoxylaceae Zeaceae Zephyranthaceae

Zingiberaceae ZiziphaceaeZosteraceae Zygophyllaceae

ON THE INTERPRETATION OF THE TREES TEXT ABBREVIATIONS ETC

For the general organisation and design of the Angiosperm Phylogeny Website see

the Introduction to the whole site Here I focus on the Phylogeny and Evolution of the

Seed Plants portion

The organization of the information here is hierarchical that is apomorphies are

mentioned only at the level at which they occur This is in line with a phylogeny- or tree-

based system Indeed if perhaps ironically a similar procedure has long been seen as an

advantage of many so-called natural systems even those that owe nothing to

evolutionary ideas (eg Cesalpino 1583 Jussieu 1789)

There is much to do to make this style of presentation fully effective As mentioned

above whether a character state that is more or less constant in a group is a

synapomorphy often awaits further clarification of relationships both within the group

and between that group and its immediate relatives For example although most

Annonaceae have stamens with distinctive prolongations of the connective if taxa

like Anaxagorea are sister to the rest of the family such connectives may not be a

synapomorphy of Annonaceae nor may indehiscent fruits and the absence of

staminodes (eg Scharaschkin amp Doyle 2005 2006) Similarly the dismemberment of

the Icacinaceae and association of fragments once in that family with Aquifoliales

Apiales and perhaps also Garryales has important effects on the characterisations of

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 51: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

51

those taxa (cf Bremer et al 2001) Gradually however characters are moving to their

appropriate levels

Remember that we know little about the functional or adaptive significance of

many synapomorphies Apart from some wind-pollinated taxa aquatics and parasites it

is usually difficult to characterise larger groups ecologically although groups like

Ericaceae are partial exceptions Furthermore much of the biology in family

discussions comes from mention of the pollination biology or other aspects of the

biology of particular genera and other small groups within a family Users of phylogeny-

based classifications such as this will add a biological emphasis as they focus on the

polination galls herbivores etc of the taxa in larger clades that grow locally However

I have been adding details concerning the diversification of clades particularly striking

associations with particular groups of herbivores or pollinators etc as I come across

them and some of this information is being summarized in the context of angiosperm

diversification as a whole in a special sectionEVOLUTION AND DIVERSIFICATION OF THE

ANGIOSPERMS In this context I am reminded of Ledyard Stebbinss remarks in his

dismissive review of the 1262 pages of Art Cronquists An Integrated System of

Classification of Flowering Plants (1981) which read in their entirety The only material

of even peripheral interest to the general evolutionist consists of short commentaries

on family relationships placed at the end of the description [sic] of many of the

families (Stebbins 1982 p 628)

The character hierarchy was built up by first drawing up lengthy descriptions of

families and then fitting the characters in the descriptions to molecular-based trees with

rather conservative topologies That is features found in characterisations of sister taxa

were removed and considered a feature of the clade that included those two taxa and

the whole process repeated The states of some characters at the base of the

angiosperm tree were fairly obvious hence the fairly lengthy characterisation

(apomorphies and plesiomorphies definitely mixed) for the angiosperms as a whole

For some of these characters I then worked up the tree placing them as high as the

evidence suggested Otherwise features in common to each clade whether order

families within an order or groups of orders are those that are as far as is known

common (reversals excepted) to all the families in that clade they may also be

synapomorphies (but see above) and are placed at the lowest level in the tree for which

I have information on the variation For some features I have used both approaches but

confusion should be minimal As relationships and our knowledge of the variation within

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 52: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

52

characters improve the top-down and bottom-up approaches merge (for more details

see the Apomorphies page)

Trees showing relationships within many orders - and within a few of the larger

families - are included Note that they may have been cobbled together from more than

one study In the trees I have emphasized mostly nodes with substantial support

(eg 80 bootstrap support) that appear after analysis of data from more than one

gene by and large there is little conflict between different studies when they overlap

Many studies now use Bayesian analysis here posterior probabilities are usually

substantially higher than bootstrap or jacknife values for the same nodes In a few cases

(eg in Santalales the base of rosids) I have been somewhat less cautious but I have

always tried to make it clear where I am treading on thin ice There are references (not

exhaustive) to papers giving support for the relationships suggested here and these

papers may have more resolved trees than those shown albeit the greater detail may

have little support Chase et al (1993 2000a) Olmstead et al (1992 1993 2000)

Olmstead and Graham (2000a b) Savolainen et al (2000a b) D Soltis et al (1997

1998 2000 2003 2005b 2007a) P Soltis et al (2000) B Bremer et al (2002) Hilu et

al (2003) etc are invaluable sources for the developing the big picture of angiosperm

relationships If you printed out all the trees here and stuck them together you might

seem to have some kind of super tree however it is clear from the description of my

modus operandi that this would hardly be a formal super tree PhyloMatic is a another

resource to be used When looking at these trees and thinking about the relationships

they suggest remember the caveats made above

As mentioned above I very largely follow the families and orders recognized by the

Angiosperm Phylogeny Group (APG 2003) with a few modifications suggested by

more recent work Families are grouped within orders as far as possible according to

their phylogenetic relationships I give some ordinal names to families that are

unassigned in APG largely for didactic purposes

For the authorities of the names of subfamilies families orders etc I have relied

heavily on Reveals listings (Reveal 2001 onwards) and especially Reveal in Thorne

(2007) These should be consulted in case of doubt since the authors of some names as

given here may be incorrect and bibliographic work that affects authority names

proceeds apace Synonymy is as complete as I can get it at the familial level and above

Most paraphyletic groups are clearly indicated as being such eg Caesalpinioideae

Olacaceae

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 53: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

53

Apomorphies are mentioned following the order in the discussion of the characters

on the Characters page You may find apparent contradictions when looking at two

nodes These usually mean that the feature mentioned more basally in the tree has

changed perhaps even reversed Thus at the node AUSTROBAILEYALES

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]] ]

you will find ethereal oils + - this part of the tree seems to be where that feature

evolved However in the characterisation of all monocots minus Acorales and of

[Ceratophyllales + eudicots] you will find ethereal oils 0 It is hypothesized that

ethereal oils were lost twice - as well as being reacquired in Zingiberaceae within

Lamiaceae etc Parentheses indicate characters that are common in a clade being

found in several but not all terminal taxa but in no obvious pattern Examples are

septal nectaries and cuticle waxes in monocots N-fixing in part of the rosid I clade and

iridoids in asterids If you are interested in seeing the apomorphies at all levels within

seed plants for a particular order you should consult the individual order pages in the

main body of the site each page starts off with a characterisation of the common

ancestor of all seed plants then of angiosperms and then characterisations of all nodes

on the branches leading up to the order in question

The contraction P stands for perianth T for tepals K for calyx C for corolla A for

the androecium as a whole and G for the gynoecium G is used most often to refer to

carpel number and if the numbers are in square brackets it means that the carpels are

connate and if underlined the ovarycarpels are superior thus G [3] means that the

gynoecium consists to three connate carpels which are superior in position Many

means that there are more than fifteen or so parts Square brackets enclose

explanations or glosses of the feature described A fuller list of abbreviations etc used

may be found underAbbreviations on the top of the left pane

Subfamilies or tribes where included are numbered sequentially within each

family Knowing something of these groupings is often important because it clarifies

which characters of families really are potential synapomorphies and which

characterise only parts of the family speciose though those parts may be - see for

example Annonaceae Convolvulaceae Fabaceae and Nyctaginaceae

Following familial and subfamilial apomorphies are two figures the approximate

number of accepted genera and species in the group I mention most genera with 50 or

more species and estimate total numbers of species and genera in families this

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 54: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

54

information is taken from Mabberleys excellent Dictionary (Mabberley 2008) accounts

in Kubitzki (1993 onwards) and Govaertss checklists (eg Govaerts amp Frodin 1998) etc

General geographical distribution is indicated and there are distribution maps for

most families on the individual order pages These maps give only approximate natural -

ie unaffected by recent human activities - distributions

Following most families and a very few orders are brief paragraphs giving

characters that I find to be helpful in recognizing the taxa the terms used here may not

be perfectly correct botanically

In the main pages there are a number of sections with subheadings following most

family accounts (and also at higher nodes in the tree when relevant) I follow the same

general sequence of sections here but without the subheadings I mention the set-up in

the main pages so this part of the Student Pages is more comprehensible The first

is Evolution Information here includes ages of clades which are being added and early

fossils comments on mycorrhizal endophytic or other fungal associations insect groups

that feed on the plants information on pollination and disseminule distribution etc

Note that clade ages at this stage of our knowledge are unreliable and in several cases

there are substantially different estimates for the same event so please treat these

dates with caution

In the paragraph Economic Importance is included only a few of the economically

globally important taxa and this part is currently notably incomplete The

section Chemistry Morphology etc summarizes interesting variation withion the

family and includes referencess to major sources of information that are not mentioned

elsewhere on the page In the Phylogenysection there are summaries of major

phylogenetic works bearing on our current ideas of phylogentic relationships in the

family In the section Classification can be found references to the infrafamilial

classification followed here and there is some discussion about generic limits in the

family and sometimes mention of important recent monographs of groups in the family

In the section Previous relationships I have included a little discussion about groupings

recognised by extant authors who have classificatory philosophies different from that

followed here Insofar as I talk about earlier ideas of relationships I mention largely

some suggestions of Cronquist (1981) and Takhtajan (1997) but only because their work

is still commonly used Note that finding out who was first in suggesting a particular

relationship is no goal of these pages the more so since what is often more interesting

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS
Page 55: APG Classification for Students

PHYLOGENY AND EVOLUTION OF SEED PLANTS 2010

55

in such situations is not that a particular suggestion was made but exactly why it was

made The section Trivia needs no explanation There is a complete family-level and

above synonymy in the approriate place on each page

Note that I have removed many of the older references that can be found on the

main pages although these may contain large amounts of invaluable comparative data

(note that much of this older literature is coming on line for example see theBotanicus

Digital Library which will soon be approaching 1000000 pages scanned) Look in the

individual order pages and especially the Characters page for references to these

older general surveys or other sources of information for particular characters

In the Glossary pages there are definitions of the terms commonly used in the site

and some other terms that may be encountered representative chemical formulae etc

Definitions as far as possible follow current usage rather than etymology or original

definitions (see eg Rickett 1954-56) Building this glossary forced me to confront head-

on such problems as the plethora of terms that have been used to describe

inflorescences and fruits - few of these are found below Note that even if we dignify

some feature of an organism by a technical term this does not mean that the term

necessarily refers to anything real about the organism Also the use of the same term

for a particular structure in two groups in no way implies that this structure is

homologous in those groups or is a synapomorphy for them or some more inclusive

group There are one or two exceptions such as carpel perhaps but these are definitely

exceptions Thus what we call stipules in Magnoliaceae and Rubiaceae monosymmetric

flowers in Orchidaceae and Fabaceae and arils in Passifloraceae and Crossosomataceae

simply fit the definitions we give to such structures - which are purely morphological

I will be more than grateful if any references in the text that lack citations and any other

errors of omission or commission are brought to my attention -

peterstevensmobotorg should find me Spelling is erratic and somewhat mid-

Atlantic grammar is little better All mistakes are mine

  • PHYLOGENY AND EVOLUTION OF SEED PLANTS