apg classification for students
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Classification of seed plants by phylogenyTRANSCRIPT
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
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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system that is followed here Books are out-of-date before they appear furthermore
there is no comprehensive phylogeny-based treatment of angiosperms out-of-date or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
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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
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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
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
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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
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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
-
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
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|Z
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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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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
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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
-
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
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|Z
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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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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
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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
-
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
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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
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|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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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
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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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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
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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
-
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
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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
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|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
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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
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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
-
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
-
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]]]]]
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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
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inghamiales Cunoniales Cupressales Cycadales Cyclanthales Cymodoceales Cynarales
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Ochnales Oenotherales Olacales Oleales Onagrales Oncothecales Opuntiales Orchid
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Balanitaceae Balanopaceae Balanophoraceae Balsameaceae Balsaminaceae Bambusa
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Colchicaceae Coleogynaceae Columelliaceae Combretaceae Commelinaceae Comocl
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aceae Cotyledonaceae CoulaceaeCoutariaceae Coutoubiaceae Crassulaceae Crescen
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aceae
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matiaceae Eupteleaceae EuryalaceaeEustrephaceae Euthemidaceae Eutocaceae Exb
ucklandiaceae Exocarpaceae
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zroyaceae FlacourtiaceaeFlagellariaceae Flindersiaceae Foetidiaceae Forestieraceae
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aliaceae Garciniaceae GardeniaceaeGardneriaceae Garryaceae Geissolomataceae Ge
itonoplesiaceae Gelsemiaceae Geniostomaceae GentianaceaeGeonomataceae Geosir
idaceae Geraniaceae Gerrardinaceae Gesneriaceae Gethyllidaceae Gilliesiaceae Gina
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onoviaceae GrossulariaceaeGrubbiaceae Guaiacanaceae Guamatelaceae Guettardac
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aptanthaceae Harmandaceae Hebenstretiaceae HectorellaceaeHederaceae Hedyosm
aceae Hedyotidaceae Hedysaraceae Heisteriaceae Heleniaceae Heliamphoraceae H
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aceae Helleboraceae Heloniadaceae HelosidaceaeHelwingiaceae Hemerocallidaceae
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teropyxidaceae HeterostylaceaeHibbertiaceae Hibiscaceae Hilleriaceae Himantandra
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Icacinaceae Idiospermaceae Illecebraceae Ilicaceae Illiciaceae Illigeraceae Impatienta
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giaceae Kirengeshomaceae KirkiaceaeKobresiaceae Koeberliniaceae Koelreuteriaceae
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aceae LimodoraceaeLimoniaceae Limosellaceae Linaceae Linariaceae Lindenbergiace
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nzieziaceae Mercurialaceae Merenderaceae Mesembryaceae Mesembryanthemaceae
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Mespilaceae MetasequioaceaeMetteniusaceae Meyeniaceae Miconiaceae Micranth
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niciaceae Phoradendraceae Phormiaceae Phrymaceae Phylicaceae Phyllanthaceae P
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hyllocladaceae PhyllonomaceaePhysenaceae Phytelephaceae Phytocrenaceae Phytol
accaceae Piceaceae Picramniaceae Picridaceae PicrodendraceaePilgerodendraceae P
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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
-
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
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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
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|Z
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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
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lismataceae Alliaceae Allioniaceae Allophylaceae Aloaceae Alopecuraceae Alpiniacea
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aceae Apiaceae ApocynaceaeApodanthaceae Aponogetonaceae Aporusaceae Apose
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raceae Aragoaceae Araliaceae Aralidiaceaee Araucariaceae Arbutaceae Arceuthidace
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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
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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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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
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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
-
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
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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
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
-
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
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|Z
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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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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
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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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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
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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
-
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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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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|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
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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
-
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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|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
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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
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
-
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
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|Z
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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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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
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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
-
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
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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
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
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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
-
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
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
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
-
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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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
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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
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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
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
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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
-
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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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
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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
-
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
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Vacciniales Vahliales Vallisneriales Velloziales Veratrales Verbenales Viburnales Vinc
ales Violales Viscales VitalesVochysiales
Welwitschiales Winterales
Xanthorrhoeales Ximeniales Xyridales
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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
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aceae Anrederaceae Anthemidaceae Anthericaceae Anthobolaceae Antidesmataceae
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aceae Apiaceae ApocynaceaeApodanthaceae Aponogetonaceae Aporusaceae Apose
ridaceae Apostasiaceae Aptandraceae AquifoliaceaeAquilariaceae Aquilegiaceae A
raceae Aragoaceae Araliaceae Aralidiaceaee Araucariaceae Arbutaceae Arceuthidace
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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
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daceae Buddlejaceae Buglossaceae Bulbocodiaceae Bumeliaceae Bupleuraceae Burc
hardiaceae BurmanniaceaeBurseraceae Butneriaceae Butomaceae Buxaceae Byblida
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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
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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
-
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
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|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
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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
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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
-
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
42
ALPHABETICAL LISTING OF ALL FAMILY NAMES OF SEED PLANTS WITH LINKS
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|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
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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
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
-
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
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
-
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
-
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
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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
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
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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
-
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
-
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
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|Z
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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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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
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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
-
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
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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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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
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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
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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
-
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
42
ALPHABETICAL LISTING OF ALL FAMILY NAMES OF SEED PLANTS WITH LINKS
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|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
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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
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
-
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
-
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
-
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
-
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
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
-
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
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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
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
-
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
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
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
-
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
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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
-
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
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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
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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
-
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
-
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
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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
-
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
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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
-
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
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
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
-
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
-
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
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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
-
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
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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
-
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
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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
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
-
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
-
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
-
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
-
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
-
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
-
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
-
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
-
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
-
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
-
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
-
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
-
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
-