an update of the angiosperm phylogeny group classiï¬cation for

Botanical Journal of the Linnean Society

2003

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399ndash436 With 1 figure

copy 2003 The Linnean Society of London


2003

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399

Blackwell Science LtdOxford UKBOJBotanical Journal of the Linnean Society 0024-4074The Linnean Society of London 2003 2003141

Original Article

CLASSIFICATION OF ORDERS AND FAMILIES OF FLOWERING PLANTSAGP II

Corresponding author Mark Chase E-mail mchaserbgkeworgukRecommended citation abbreviated as lsquoAPG II 2003rsquo This paper was compiled by Birgitta Bremer Karingre Bremer Mark W ChaseJames L Reveal Douglas E Soltis Pamela S Soltis and Peter F Stevens who were equally responsible and listed here inalphabetical order only with contributions from Arne A Anderberg Michael F Fay Peter Goldblatt Walter S Judd MariKaumlllersjouml Jesper Karingrehed Kathleen A Kron Johannes Lundberg Daniel L Nickrent Richard G Olmstead Bengt Oxelman JChris Pires James E Rodman Paula J Rudall Vincent Savolainen Kenneth J Sytsma Michelle van der Bank KennethWurdack Jenny Q-Y Xiang and Sue Zmarzty (in alphabetical order) Addresses B Bremer The Bergius Foundation at the RoyalSwedish Academy of Sciences PO Box 50017 SE-104 05 Stockholm Sweden K Bremer Department of Systematic BotanyEvolutionary Biology Centre Uppsala University Norbyv 18D SE-752 36 Uppsala Sweden M W Chase Jodrell LaboratoryRoyal Botanic Gardens Kew Richmond Surrey TW9 3DS UK J L Reveal University of Maryland 18625 Spring Canyon RoadMontrose Colorado 81401ndash7906 USA D E Soltis Department of Botany University of Florida Gainesville Florida 32611ndash8526 USA P S Soltis Florida Museum of Natural History Dickinson Hall University of Florida Gainesville Florida 32611ndash7800 USA and P F Stevens Department of Biology University of Missouri-St Louis and Missouri Botanical Garden PO Box299 St Louis Missouri 63166ndash0299 USA

An update of the Angiosperm Phylogeny Group classification for the orders and families offlowering plants APG II

THE ANGIOSPERM PHYLOGENY GROUP

Received June 2002 accepted for publication December 2002

A revised and updated classification for the families of the flowering plants is provided Newly adopted orders includeAustrobaileyales Canellales Gunnerales Crossosomatales and Celastrales Pertinent literature published since thefirst APG classification is included such that many additional families are now placed in the phylogenetic schemeAmong these are Hydnoraceae (Piperales) Nartheciaceae (Dioscoreales) Corsiaceae (Liliales) Triuridaceae (Pan-danales) Hanguanaceae (Commelinales) Bromeliacae Mayacaceae and Rapateaceae (all Poales) Barbeuiaceae andGisekiaceae (both Caryophyllales) Geissolomataceae Strasburgeriaceae and Vitaceae (unplaced to order butincluded in the rosids) Zygophyllaceae (unplaced to order but included in eurosids I) Bonnetiaceae Ctenolopho-naceae Elatinaceae Ixonanthaceae Lophopyxidaceae Podostemaceae (Malpighiales) Paracryphiaceae (unplaced ineuasterid II) Sladeniaceae Pentaphylacaceae (Ericales) and Cardiopteridaceae (Aquifoliales) Several major fami-lies are recircumscribed Salicaceae are expanded to include a large part of Flacourtiaceae including the type genusof that family another portion of former Flacourtiaceae is assigned to an expanded circumscription of AchariaceaeEuphorbiaceae are restricted to the uniovulate subfamilies Phyllanthoideae are recognized as Phyllanthaceae andOldfieldioideae as Picrodendraceae Scrophulariaceae are recircumscribed to include Buddlejaceae and Myoporaceaeand exclude several former members these are assigned to Calceolariaceae Orobanchaceae and Plantaginaceae Weexpand the use of bracketing families that could be included optionally in broader circumscriptions with otherrelated families these include Agapanthaceae and Amaryllidaceae in Alliaceae

sl

Agavaceae Hyacinthaceae andRuscaceae (among many other Asparagales) in Asparagaceae

sl

Dichapetalaceae in Chrysobalanaceae Turner-aceae in Passifloraceae Erythroxylaceae in Rhizophoraceae and Diervillaceae Dipsacaceae Linnaeaceae Mori-naceae and Valerianaceae in Caprifoliaceae

sl


Botanical Journal of theLinnean Society

2003

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399ndash436

ADDITIONAL KEYWORDS

angiosperms ndash gene sequences ndash phylogenetics

INTRODUCTION

During the 1990s reconstruction of flowering plantphylogeny took a great step forward Rapidly accumu-

lating DNA sequences in particular from the plastidgene

rbcL

(eg Chase

et al

1993) provided new andinformative sets of data Cladistic analysis of thesedata sets was also much improved especially through

400

AGP II



2003

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399ndash436

development of phylogenetic theory and application toanalysis of large data sets (eg Hillis 1996) and var-ious methods for estimating the support for individualclades in the phylogenetic trees (Felsenstein 1985Farris

et al

1996) The outline of a phylogenetic treeof all flowering plants became established and severalwell supported major clades involving many familiesof flowering plants were identified In many cases thenew knowledge of phylogeny revealed relationships inconflict with the then widely used modern classifica-tions (eg Cronquist 1981 Thorne 1992 Takhtajan1997) which were based on selected similarities anddifferences in morphology rather than cladistic anal-ysis of larger data sets involving DNA sequences orother forms of systematic data It became clear thatnone of the previous classifications accuratelyreflected phylogenetic relationships of floweringplants and communication about plant evolutionreferring to the old classification schemes becameincreasingly difficult To alleviate this problem agroup of flowering plant systematists calling them-selves the Angiosperm Phylogeny Group (APG forshort) proposed a new classification for the families offlowering plants (APG 1998)

The initial APG (1998) system comprised 462 fami-lies arranged in 40 putatively monophyletic ordersand a few monophyletic higher groups The latter werenamed informally as monocots commelinoids (herechanged to commelinids to prevent confusion withsubfamily Commelinoideae of Commelinaceae) eud-icots core eudicots rosids including eurosids I and IIand asterids including euasterids I and II The focuswas on orders and less on families An attempt wasmade to recognize orders well supported as monophyl-etic in large jackknife analyses of molecular data(Kaumlllersjouml

et al

1998) In general the orders werefairly widely circumscribed especially in comparisonwith those of Takhtajan (1997) A few monofamilialorders were recognized (Ceratophyllales Acorales andArecales) for cases in which these families were appar-ently sister groups of larger clades including severalorders Many families were not classified to orderbecause their positions were uncertain or unknownand these families were listed under the supraordinalgroups where they were known to belong or at the endof the system in a list of families probably eudicots ofuncertain position APG predicted that there would belittle need to change the circumscription of the ordersexcept for inclusion of families not then assigned toorder and possible transfer of occasional misplacedfamilies It was also realized that new orders might beestablished if monophyletic groups of unplaced fami-lies were identified

The APG system also involved the recognition ofstrictly monophyletic groups at all levels but it wasacknowledged that there were families known to be

non-monophyletic (eg Euphorbiaceae and Scrophu-lariaceae) Reclassification of these into monophyleticunits was not possible in 1998 and required furtherinvestigation Furthermore monophyly of many fam-ilies remained to be investigated with extensive sam-pling and application of molecular phylogenetictechniques Thus it was acknowledged that somechanges in family circumscription would be necessaryto reflect improved understanding of phylogeneticrelationships For some families already investigatedand found to be monophyletic alternative optionalcircumscriptions were indicated by listing the sisterfamily or families in square brackets immediatelyafter the family For example Nymphaeaceae could beinterpreted either to exclude or include a sister familyCabombaceae

Five years have now passed since the APG systemwas compiled Recent advances in our knowledge offlowering plant phylogeny indeed have motivated sev-eral changes in family circumscription and classifica-tion as well as the addition of a few new orders Wetherefore present here an updated version of the APGsystem

In general we have adopted a conservativeapproach and propose here changes in the APG sys-tem only when there is substantial new evidence sup-porting a revised classification Five additional ordersare recognized Austrobaileyales Canellales Celas-trales Crossosomatales and Gunnerales These rep-resent well-supported monophyletic groups offamilies unclassified to order in APG (1998) Circum-scription of none of the APG orders has been changedexcept for the addition of a number of the familiesunclassified to order in APG (1998) When morerecent analyses have demonstrated that such familiesof formerly uncertain position are well nested insidethe APG orders or well supported as sister groups toany of the APG orders the latter have been expandedto include these families Thus some APG ordershave been more widely circumscribed to include theirsister groups (eg Adoxaceae being included in Dipsa-cales cf Bremer 2000) except in one case in whichthe pair of Canellaceae and Winteraceae has beenestablished as an order Canellales rather thanincluded in their sister group Piperales (mostresearchers would consider these two groups toodivergent to include in a single order) No APG ordershave been merged or split and no families have beentransferred from one order to another Only in onecase has a family been removed from an APG orderOncothecaceae have been excluded from Garryalesand assigned to a position at the beginning of theeuasterids I without classification to order becauserecent analyses have not supported any clear (iebootstrap- or jackknife-supported) ordinal position forthat family

CLASSIFICATION OF ORDERS AND FAMILIES OF FLOWERING PLANTS

401



2003

141

399ndash436

Figure 1

Interrelationships of the orders and some families supported by jackknife or bootstrap frequencies above 50in large-scale analyses of angiosperms All except five of the clades are supported by the Soltis

et al

(2000) analysis of18S rDNA

rbcL

and

atpB

sequences from a wide sample of angiosperms Three clades Canellales

+

PiperalesLaurales

+

Magnoliales and these four orders together are supported by analyses of several different gene sequences ofphylogenetically basal angiosperms (Qiu

et al

1999 Graham amp Olmstead 2000) One clade that of all core eudicotsexcept Gunnerales is supported by analysis of

rbcL

sequences from a wide sample of eudicots (Savolainen

et al

2000)Another clade that of all asterids except Cornales is supported by a six-marker analysis of a wide sample of asterids(Bremer

et al

2002) Rosid and asterid families not classifed to order are not shown

402

AGP II



2003

141

399ndash436

Interrelationships among the orders and some ofthe unclassified families are now better understoodthan they were when the APG system was developedIn Figure 1 we show interrelationships of the ordersand some families supported by jackknife or bootstrappercentages above 50 in large-scale analyses of 18SrDNA

rbcL

and

atpB

sequences from a wide sampleof angiosperms (Soltis

et al

2000a) Similar relation-ships were shown based on analyses of several genesfrom phylogenetically basal angiosperms (Qiu

et al

1999 Graham amp Olmstead 2000 Zanis

et al

2002)and of

rbcL

sequences with a wide sample of eudicots(Savolainen

et al

2000b cf also Kaumlllersjouml

et al

1998) However relationships among the major ordersof monocots and core eudicots and partly among theorders of rosids and asterids are still uncertain(Fig 1 Chase

et al

2000 Soltis

et al

2000a)At the family level several families have been syn-

onymised or re-circumscribed especially in Aspar-agales Malpighiales and Lamiales A few familieshave been re-established from synonymy to makethem monophyletic in so far as it is possible given cur-rent knowledge about generic interrelationships As ageneral principle we have avoided changing circum-scription of the families unless necessary to preservemonophyly There are however two exceptions to thisgeneral principle of stability First detailed workwithin some taxa since APG (1998) has generatedmuch new knowledge about interrelationships andwhen specialists have proposed a new and well sup-ported classification it has been followed even if ourprevious classification did comprise monophyleticfamilies Second in several cases accumulating knowl-edge of phylogeny has demonstrated sister-group rela-tionships involving small monogeneric families Suchtaxa represent redundancies in classification andhence we have usually reduced monogeneric familiesto synonymy to reduce this redundancy In some caseshowever we have retained the existing family classi-fication when it was judged that a monogeneric familyis so different morphologically from its sister groupthat merging the two would create a morphologicallyunrecognizable entity We recognize that decisionsusing the argument lsquotoo divergent morphologicallyrsquoare likely to be highly subjective and largely intuitivebut these arguments are a long established traditionWe generally accept the opinion of specialists in suchcases but we also recognize that specialists nearlyalways favour splitting of groups they view as lsquotoo het-erogeneousrsquo In several cases we have listed familiesin brackets indicating the possibility of alternativecircumscriptions as described in the introduction tothe APG system above With the changes introducedhere the number of orders has increased from 40 to 45and the number of families decreased from 462 to 457Of this number 55 families are listed in brackets We

are aware of at least one appropriate additional fam-ily that has yet to be formally proposed Summarizedbelow are the changes to APG (1998) with appropriatereferences supporting these alterations Since 1998five proposed systems for the angiosperms have beenpublished Two (Judd

et al

1999 2002 Stevens 2001)more or less follow the system presented in APG(1998) One (Thorne 2001 pers comm) approachesthat of APG whereas two others (Doweld 2001Wu

et al

2002) basically follow that proposed byTakhtajan (1997)

ROOT OF THE ANGIOSPERM TREEAND MAGNOLIIDS

Relationships at the base of the angiosperms havebeen largely clarified with most analyses supporting

Amborella

as sister to all other extant angiosperms(Qiu

et al

1999 Soltis

et al

2000a) In contrast tothese studies Barkman

et al

(2000) found strong sup-port for Nymphaeaceae

Amborella

as sister to all otherangiosperms in an analysis from which lsquonoisy datarsquowere removed Further analyses by Zanis

et al

(2002)rejected the Nymphaeaceae

Amborella

rooting nearlyall tests found strong support for

Amborella

alone assister to the rest with Nymphaeaceae as the subse-quent sister to the rest Either one order with bothfamilies or two orders might still be possible so werefrain from formally proposing names for these theordinal names Amborellales and Nymphaeales areavailable Austrobaileyales are recognized by APG forthe first time and comprise Austrobaileyaceae Trime-niaceae and Schisandraceae (optionally includingIlliciaceae) A clade of

Austrobaileya

Illicium

and

Schisandra

received 99 jackknife support in analy-ses of

rbcL

atpB

and 18S rDNA (Soltis Soltis amp Chase1999 Soltis

et al

2000b) Material of

Trimenia

wasnot available for these multigene analyses howeverparsimony analyses of

rbcL

(Renner 1999) and 26SrDNA (Soltis

et al

2000b) and a maximum likelihoodanalysis of

rbcL

atpB

and 18S rDNA (Soltis

et al

2000b) for fewer taxa placed

Trimeni

a in this cladeBootstrap support for this clade in 5- 6- and 11-geneanalyses was 100 (Qiu

et al

1999 Zanis

et al

2003)The magnoliids a superordinal group comprise

Laurales Magnoliales Piperales and a new APGorder Canellales with two families Canellaceae andWinteraceae This larger magnoliid clade did notreceive jackknife support greater than 50 in thethree-gene analyses of Soltis

et al

(1999) and Soltis

et al

(2000a) but with the addition of more genesbootstrap support for this clade increased to 64(Zanis

et al

2003) and 67 (Qiu

et al

1999) for fivegenes and 100 in a compartmentalized analysis ofsix genes (Zanis

et al

2002) and 11 genes (Zanis

et al

2003) Within the magnoliids Laurales and Magno-


403



2003

141

399ndash436

liales are sisters (71 bootstrap support Qiu

et al

1999 100 Zanis

et al

2003) and Piperales andCanellales are sisters (83 bootstrap support Qiu

et al

1999 100 Zanis

et al

2003) However anal-ysis of 104 morphological characters for 52angiosperms (Doyle amp Endress 2000) resulted in dif-ferent relationships among these clades Magnoliales

+

Canellales were sister to Laurales and Piperaleswere distantly related in a polytomy with the mono-cots Nymphaeaceae and several clades of eudicotsNone of these relationships received bootstrap supportgreater than 50

The sister-group relationship of Winteraceae andCanellaceae has received bootstrap or jackknife sup-port of 99 or 100 in all recent multigene analyses(eg Soltis

et al

1999 Qiu

et al

1999 Soltis

et al

1999 Zanis

et al

2002 2003) Doyle amp Endressrsquo(2000) morphological analysis also found this sistergroup (bootstrap support

lt

50) Their sister groupPiperales consists of Aristolochiaceae LactoridaceaePiperaceae and Saururaceae (APG 1998) to whichwe now add Hydnoraceae (Nickrent

et al

2001)Although the exact placement of Hydnoraceae withinPiperales is uncertain it clearly falls within this clade(Nickrent amp Duff 1996 Nickrent

et al

1998 2001) Inrecent analyses

Lactoris

appears within a clade ofAristolochiaceae as sister to

Aristolochia

+

Thottea

(Qiu

et al

1999 Zanis

et al

2003) or

Aristolochia

alone (Soltis

et al

2000a)

Thottea

was not included inthe last analysis but support for the embedded posi-tion of

Lactoris

was weak (66 or less) even with fivegenes Morphological analyses likewise recognize aclade of Piperaceae Saururaceae Aristolochiaceaeand

Lactoris

Given the uncertain position of

Lactoris

in both molecular and morphological trees we recom-mend that Lactoridaceae be retained until more con-vincing evidence of placement is obtained

The position of Chloranthaceae also requires furtherstudy It is sister to the magnoliids

+

eudicots in thesix-gene compartmentalized analysis (84 bootstrapsupport Zanis

et al

2003) but this is the only anal-ysis that has provided support for the placement of thisfamily At this time we prefer not to assign Chloran-thaceae to an order until its position becomes clearerThe name Chloranthales is available should Chloran-thaceae require assignment of a name at that rank

MONOCOTS

Although the sister group of the monocots remainsunclear a great deal of progress has been made withinthe monocots since the last APG installment Chase

et al

(2000) published a review of relationships and aproposed revision of the APG system for the monocotsbut nonetheless we will here provide information onchanges since APG (1998) The rooting of the monocots

between Acoraceae (Acorales) and the rest continuesto be supported The Chase

et al

(2000) and Soltis

et al

(2000a) analyses of three genes both agreed onthis placement as did that of Fuse amp Tamura (2000)which examined a relatively small number of mono-cots with plastid

matK

sequences Of the five familiesunplaced in APG (1998) we now have clear placementof all but Petrosaviaceae (which now also includes

Japanolirion

) The name Petrosaviales is available forthe family if it is assigned to its own order Triuri-daceae are placed in Pandanales probably as sister toPandanaceae and Corsiaceae are included in Liliales(Neyland 2002) Within Dioscoreales several changesare made as a result of the extensive study of the orderby Caddick

et al

(2000 2002ab) which used an anal-ysis of three genes

rbcL atpB

and 18S rDNA andmorphology to examine relationships of nearly all gen-era of the order Thismiaceae are sister to Burmanni-aceae which makes it reasonable to include themtogether

Trichopus (formerly Trichopodaceae) is sisterto Avetra (Dioscoreaceae) and this pair is sister toTacca (Taccaceae) All other genera of Dioscoreaceae(Rajania Nonarapenta Tamus etc) are embedded inDioscorea so a simplified taxonomy of Dioscoreaceaewould be to include these in Dioscorea and eliminateTaccaceae and Trichopodaceae (both monogeneric) byincluding them in an expanded DioscoreaceaeAlthough bootstrap support is not exceptionally higha position for Nartheciaceae in Dioscoreales is congru-ent with the non-DNA analyses of Caddick et al(2000)

Continued work on Asparagales (Fay et al 2000b)clarified relationships within the order In recentyears new families were published to accommodategenera that fell as sister taxa to clades composed ofseveral families sensu Dahlgren Clifford amp Yeo (1985)but this process has led to both a rearrangement offamily limits and an increased recognition of monoge-neric and small families Specialists in these familieshave hoped to take a broader view of family limits inAsparagales which is now possible because the pat-terns are relatively clear (Fay et al 2000b) BecauseDahlgren and co-workers believed that broadly cir-cumscribed concepts of Liliaceae were grossly unnat-ural they recognized as families only those groups inwhich they had some confidence of monophyly Theresult of this approach was circumscription of nar-rowly defined families When molecular systematiststurned their attention to relationships of the lilioidmonocots (Duvall et al 1993 Chase et al 1995ab)they used this classification as the basis of their sam-pling Hence they retained the circumscriptions ofDahlgren et al (1985) without further consideration ofwhether these units should in fact be recognized asfamilies APG (1998) also used this system and soAsparagales was established with 29 recognized fam-

404 AGP II

copy 2003 The Linnean Society of London Botanical Journal of the Linnean Society 2003 141 399ndash436

ilies many of them small (1ndash5 genera) Although thisapproach seemed logical at the time it now in retro-spect appears less so because it bequeathed usextremely narrowly defined family limits Only spe-cialists in this group understand this taxonomy and itis so difficult to teach that many instructors use Lili-aceae sl rather than the APG system We thereforepropose here to simplify the higher Asparagales byoptionally reducing the number of families to two Alli-aceae and Asparagaceae These can easily be identi-fied by the umbellate inflorescences of Alliaceae(optionally including both Agapanthaceae and Ama-ryllidaceae) vs the racemes of Asparagaceae exceptfor Themidaceae with umbels but these have manysubtending and internal bracts whereas those of Alli-aceae typically have just two (or if there are more theyare not internal in the umbel Pires amp Sytsma 2002)In Asparagaceae we optionally include AgavaceaeAnemarrhenaceae (monogeneric) AnthericaceaeAphyllanthaceae (monogeneric) Behniaceae (monoge-neric) Herreriaceae (two genera) HyacinthaceaeLaxmanniaceae Ruscaceae and Themidaceae InRuscaceae Rudall Conran amp Chase (2000a) alreadyincluded Convallariaceae Dracaenaceae (three gen-era) Eriospermaceae (monogeneric) and Nolinaceae(2ndash3 genera) We propose here to use the bracketingsystem to indicate that those who wish to recognizesome additional monophyletic groups may continue todo so and still use the lsquoAPG systemrsquo However in thatcase we would recommend that Agavaceae shouldinclude Anemarrhenaceae Anthericaeae Behniaceaeand Herreriaceae (these are listed in the family syn-onymy in the appendix) Along the same lines we listXanthorrhoeaceae sl as optionally including bothAsphodelaceae and Hemerocallidaceae (which alreadyincluded Phormiaceae of earlier authors) We realizethat some researchers may be perturbed by this fur-ther re-organization of family lines within Aspar-agales but we believe this modification provides amuch-needed simplification of familial taxonomy inthis order

We were prompted to make the changes to Aspar-agales taxonomy by the condensation of families thathas already been made in Liliales Relative to the sys-tem of Dahlgren et al (1985) APG (1998) had alreadyreduced Calochortaceae Petermanniaceae Trilli-aceae Tricyrtidaceae and Uvulariaceae and we mayyet include Philesiaceae and Rhipogonaceae in Smila-caceae (following previous authors on account of theirspinose pollen Rudall et al 2000b) At this time theonly change we make is the addition of the myco-parasitic Corsiaceae on the basis of 26S rDNA data(Neyland 2002) Pandanales have the same circum-scription except for the addition of another achloro-phyllous family Triuridaceae based on analyses of18S rDNA sequence data (Chase et al 2000)

In the commelinids (we change the name here toavoid confusion with Commelinoideae) the relation-ships of many of the previously unplaced families havebeen resolved (as summarized in Chase et al 2000)Abolbodaceae are included in Xyridaceae in whichmost recent treatments have placed them Bromeli-aceae Mayacaceae and Rapateaceae are all includedin Poales and Hanguanaceae are moved to Commeli-nales Poales are now a large order of 18 families andsome researchers have advocated splitting them intoas many as three or four orders (Givnish et al 1999Judd et al 1999 Thorne 2001 pers comm) but untilrelationships are more clearly resolved such demoli-tion would be premature We also point out thatAsparagales as circumscribed here is still larger andmore diverse morphologically Based on Chase et al(2000) there is clear evidence that Poales are mono-phyletic but some relationships within the orderremain unclear Bremer (2002) analysed family inter-relationships within Poales using combined rbcLatpBanalyses and found strong support for cyperoid(Cyperaceae Juncaceae and Thurniaceae) and grami-noid clades (Anarthriaceae Centrolepidaceae Ecdeio-coleaceae Flagellariaceae Joinvilleacae Poaceae andRestionaceae) Within the latter clade Ecdeio-coleaceae rather than Joinvilleacae were found to besister to Poaceae Although the two large clades aboveare now clearly defined their relationships to theother families of Poales requires further work

We have not adopted the new monogeneric familiescarved out of Anarthriaceae (Briggs amp Johnson 2000)simply because they are monogeneric and clearlyrelated to Anarthriaceae notwithstanding theargument made by the authors that they share fewmorphological characters with each other and Anar-thriaceae The sole remaining unplaced commelinidfamily is Dasypogonaceae for which the ordinal nameDasypogonales is available should recognition becomeappropriate

Monocot phylogenetics have made immense stridesover the past 8 years due primarily to the foci providedby the two international monocot symposia held in1993 and 1998 (at the Royal Botanic Gardens KewRudall et al 1995 and the Royal Botanic GardensSydney Wilson amp Morrison 2000 respectively) Thesemeetings have focused attention both on what wasknown and more importantly on which groupsneeded additional attention As a result we now knowmore about monocots than any other major group ofangiosperms a situation that is a remarkable achieve-ment given the paucity of information available in1985 (Dahlgren et al 1985) This model should now beadopted for the other large groups of angiosperms sothat attention is likewise focused on integration ofresearch programmes and gaps in the data base Eventhe relatively well-studied asterid orders have new

CLASSIFICATION OF ORDERS AND FAMILIES OF FLOWERING PLANTS 405


members that desperately need integration into theoverall picture of eudicot evolution

EUDICOTS

Relatively few changes have been made among thefamiliesorders forming a grade at the base of the eud-icots We have placed Didymelaceae as an optionalsynonym of Buxaceae and Buxales is available if Bux-aceae were to be elevated to ordinal status Sabiaceaeand Trochodendraceae likewise remain unplaced toorder but if either or both of these changes becomesappropriate Sabiales and Trochodendrales have pre-viously been published Proteales remain unchangedexcept that we have indicated that Platanus option-ally could be included in Proteaceae although manybotanists in both Northern and Southern Hemi-spheres will probably object to this change for twotaxa that have never before been associated Ranun-culales remain unchanged from APG (1998)

Aextoxicaceae are clearly closely related to Berberi-dopsidaceae (Soltis et al 2000a among several) andthese two small families (one and two genera respec-tively) as yet have no clear relationship to the othereudicot orders so we continue to leave them unplacedto order If an ordinal name should be required (egSoltis et al 2003) Berberidopsidales is available (seebelow) It is unclear on what morphological grounds amerger of these two families could be justified thesegenera are remarkably divergent considering the sim-ilarity of their DNA sequences

Dilleniaceae were consistently placed as sister toCaryophyllales in the three-gene analysis of Soltiset al (2000a) but with jackknife support of only 60and on this basis we refrain from adding them toCaryophyllales Although the name Dilleniales isavailable it would be against the philosophy of APG tocreate a monofamilial order for them if they werefound to have a clear relationship to another recog-nized order in this case Caryophyllales

Relationships in Caryophyllales continue to be in astate of flux and therefore difficult to discuss Apartfrom Rhabdodendraceae there seem to be two majorlineages The first is composed of Caryophyllales intheir long-standing restricted sense plus Simmonds-iaceae and Asteropeiaceae + Physenaceae as succes-sive sister groups to the core members The secondincludes Ancistrocladaceae and their mostly carnivo-rous relatives (Meimberg et al 2000 Cueacutenoud et al2002) Tamaricaceae + Frankeniaceae and Plumbagi-nacae + Polygonaceae (Kaumlllersjouml et al 1998 Soltiset al 2000a Cueacutenoud et al 2002) Unfortunately thenew members of the first lineage (AsteropeiaceaePhysenaceae and Simmondsiaceae) are poorly stud-ied and some features that make the core familiesappear distinctive need re-evaluating from the per-

spective of their new alignment Within the coregroup relationships remain uncertain Appelquist ampWallace (2000) and Cueacutenoud et al (2002) found thatthe distinctive Madagascaran Didiereaceae aresister to Calyptrotheca of Portulacaceae HoweverDidiereaceae are not yet reduced to synonymy underPortulacaceae Furthermore Cueacutenoud et al (2002)found that there is a well supported but internallyunresolved group the lsquosucculentrsquo clade of Manhart ampRettig (1994) that includes Basellacaeae CactaceaeDidiereaceae Halophytaceae and PortulacaceaeAlthough Portulacaceae are clearly paraphyletic ascurrently circumscribed the composition and relation-ships of the lineages within Portulacaceae need fur-ther study before taxonomic realignment begins(hence the lack of change in the classification)

Within one of the other major clades of the coreCaryophyllales a similar problem to that of theapparently polyphyletic Portulacaceae is encoun-tered Phytolaccaceae are grossly polyphyletic rela-tive to Aizoaceae Nyctaginaceae and SarcobataceaeWe have recognized here Barbeuiaceae and Giseki-aceae both are well supported as excluded from Phy-tolaccaceae and are resurrected from the list offamilial synonyms in APG (1998) Lophiocarpus isalso clearly unrelated to the PhytolaccoideaeRivi-noideae clade but it has never been recognized as afamily (the name proposed by Bortenschlager 1973is not validly published) Corbichonia (usually Mol-luginaceae) is sister to Lophiocarpus and the pair iswell removed from the rest of Molluginaceae(Cueacutenoud et al 2002) The third major clade of corecaryophyllids is unproblematic and includes Achato-carpaceae Amaranthaceae and CaryophyllaceaeRelationships and taxonomy of the other major cladeof Caryophyllales remain as they were in APG (1998)Although additional genera and new data have beenadded no new patterns for general relationshipshave emerged (Cueacutenoud et al 2002)

Relative to APG (1998) no changes to the composi-tion in Santalales have been made (see Nickrent ampMaleacutecot 2001 and Nickrent 2002 for a summary ofrelationships) At least one of the families recognizedOlacaceae is problematic and ongoing studies ofgeneric relationships should provide evidence of howto realign family limits (Nickrent 2002) In all short-est trees produced in the combined analysis of threegenes by Soltis et al (2000a) Santalales were the sis-ter group of Dilleniaceae + Caryophyllales but withless than 50 jackknife support If they were in thefuture to receive strong support as sister to this cladethey would nonetheless be maintained because theAPG philosophy is not to alter ordinal recognitionexcept to add additional ones as needed for groupsdemonstrated to be sister to clades composed of sev-eral orders

406 AGP II


The composition of Saxifragales is one of the majorsurprises of molecular phylogenetic analyses of theangiosperms (Chase et al 1993 Morgan amp Soltis1993 Soltis et al 1997 Soltis amp Soltis 1997 Qiuet al 1998 Hoot Magallon-Puebla amp Crane 1999Savolainen et al 2000a Soltis et al 2000a) Thiseclectic assemblage comprises taxa placed in threesubclasses in modern classifications (eg Cronquist1981 Takhtajan 1997) Several changes are sug-gested here compared to APG (1998)

Phylogenetic analyses of a five-gene data set forSaxifragales (c 9000 bptaxon) (Fishbein Hufford ampSoltis 2003) have identified several major well-supported clades There is strong support for a clade ofSaxifragaceae and several woody members of theformer Saxifragaceae sensu Engler (1930 ie the cur-rently recognized families Grossulariaceae Iteaceaeand Pterostemonaceae) Within this clade the sister-group relationship between Iteaceae and Pterostemo-naceae is strongly supported Consideration should begiven to reducing Pterostemonaceae to Iteaceae byadding Pterostemon (two species) to that family A sec-ond strongly supported clade includes Crassulaceaeas sister to a clade of Haloragaceae Tetracarpaea(Tetracarpaeaceae) Penthorum (Penthoraceae) andAphanopetalum (formerly of Cunoniaceae) all smallgenera that could be combined to form a singleexpanded family Haloragaceae (Fishbein et al 2003)

Although the composition of Saxifragales nowappears clear the position of the clade among the coreeudicots is uncertain The placement of the order hasvaried among the broad phylogenetic analyses con-ducted to date Initial analyses of rbcL sequences(Chase et al 1993) placed the order as sister to allother rosids whereas analyses of atpB sequencesplaced the clade as sister to a large clade containingmost of the core eudicots (Savolainen et al 2000a)None of these placements received jackknifebootstrapsupport gt50 The three-gene analysis (Soltis et al1999 Soltis et al 2000a) placed Saxifragales as sisterto the rosids but with only weak jackknife support(60) Analyses of a four-gene data set for eudicotsindicated placement of Saxifragales as sister to allother core eudicots except Gunnerales (Soltis et al2003)

ROSIDS

Our knowledge of the composition of and relationshipsamong the rosid and eurosid I taxa has improved sig-nificantly particularly within Malpighiales and weprovide changes to reflect these newly understoodrelationships Changes to the classification elsewherein the rosids are few Geissolomataceae and Strasbur-geriaceae previously unplaced and Vitaceae previ-ously an unplaced core eudicot are added to the

rosids Vitaceae may be sister to the rest of the rosids(Soltis et al 2000a) but jackknife support for thisposition was only moderate Crossosomatales newlycircumscribed and recognized here include Crossoso-mataceae (Sosa amp Chase 2003) Stachyuraceae andStaphyleaceae all previously unplaced rosids (Soltiset al 1999 2000a Nandi Chase amp Endress 1998Savolainen et al 2000a) Crossosomatales share aseed character in which the cell walls of the many-layered testa are all or mostly lignified Seed anatomycontinues to be a valuable source of new systematicinformation that is highly congruent with phyloge-netic relationships inferred from analyses of molecu-lar data (see Doweld 2001) Circumscription of thisorder is conservative other unassigned rosid generaoften recognized as families (eg Geissoloma Ixerbaand Strasburgeria) have similar testa anatomy andmay be added to this order if support for this broadercircumscription strengthens

In Geraniales there is abundant morphological andmolecular evidence indicating that the small familiesFrancoaceae Greyiaceae and Melianthaceae areclosely related (Ronse Decraene amp Smets 1999Savolainen et al 2000b) Greyiaceae are here syn-onymised under Melianthaceae with Francoaceae anoptional further synonym Likewise Hypseocharita-ceae are an optional synonym of Geraniaceae as inAPG (1998)

In Myrtales recent work (Conti Litt amp Sytsma1996 Conti Baum amp Sytsma 1999) confirmed familycircumscriptions Clausing amp Renner (2001) showed awell-supported sister-group relationship betweenMelastomataceae and Memecylaceae clarifying thecircumscriptions of both families the two have beencombined before (eg Cronquist 1981) and havingthis option seems reasonable (they are thereforebracketed in the classification)

Zygophyllaceae and Krameriaceae are now includedin eurosid I (Soltis et al 2000a Savolainen et al2000a) Krameriaceae (monogeneric) can be includedin the already heterogeneous Zygophyllaceae (for thelatter see Sheahan amp Chase 2000) but Krameriashares few traits that could be considered synapomor-phies with Zygophyllaceae However some research-ers (eg Sheahan and Chase pers comm) see littleadvantage in the maintenance of a monogeneric fam-ily with a clear relationship to another regardless ofhow divergent the genus is from the others included IfZygophyllaceae continue to be placed as sister to aclade composed of several orders and ordinal status isappropriate then the name Zygophyllales is available

Several of the previously unplaced eurosid I familiesare now combined with Lepidobotryaceae and Celas-traceae in a newly accepted order Celastrales (Nandiet al 1998 Savolainen et al 2000b) although thegroup is not easy to characterize morphologically



Huaceae have sometimes appeared with this clade(Soltis et al 2000a) but without enough support orconsistency to warrant their inclusion here Stackhou-siaceae kept separate in APG (1998) are now syn-onymised with Celastraceae (Savolainen et al 2000aSimmons et al 2001)

The circumscription of the nitrogen-fixing clade andthe composition of the four orders included thereFabales Rosales Cucurbitales and Fagales remainlargely unchanged (see also Savolainen Spichiger ampManen 1997 Jeong Ritchie amp Myrold 1999) Rela-tionships within Rosales and especially within theCannabaceae - Cecropiaceae - Celtidaceae - Moraceae -Ulmaceae-Urticaceae complex have been problem-atic Celtidaceae are paraphyletic and includeCannabaceae and Cecropiaceae are embedded withinUrticaceae (Ueda Kosuge amp Tobe 1997 WiegrefeSytsma amp Guries 1998 Sytsma et al 2002) and it istherefore appropriate to recognize altered circum-scriptions of these families within the urticalean com-plex Within Fagales monogeneric Rhoipteleaceae arestrongly supported as sister to Juglandaceae and sothe option of combining the two is offered Howeverthe two differ considerably in their gynoecia andovules

Changes in Malpighiales mainly reflect assignmentto this order of six previously unplaced families andthe dismemberment of broadly circumscribed Flacour-tiaceae and Euphorbiaceae Of the families assignedto Malpighiales since APG (1998) Bonnetiaceae andElatinaceae have a distinctive exotegmen similar tothat of Clusiaceae and Bonnetiaceae and Clusiaceaeshare distinctive xanthones Xanthones are alsoreported from some Podostemaceae (in which Tris-tichaceae previously an unplaced rosid now areincluded) and both tenuinucellate ovules and exudateare known from Clusiaceae as well as at least somePodostemaceae (eg Contreras Scogin amp Philbrick1993 Jaumlger-Zuumlrn 1997) Relationships within theClusiaceae-Bonnetiaceae-Podostemaceae clade arehowever still unclear Ploiarium (Bonnetiaceae) hasbeen included in Malvales (Savolainen et al 2000a)but this is likely to be based on misidentified ma-terial (M W Chase pers comm) NeverthelessPodostemaceae for which the exact relationship withother angiosperms has long been controversial (Cussetamp Cusset 1988 and references therein) are finallyclose to finding a phylogenetic home Other familiesassigned to Malpighiales include CtenolophonaceaeIxonanthaceae Peridiscaceae and Lophopyxidaceae(Savolainen et al 2000a)

Recent work has clarified the limits of sets of generapreviously assigned to Flacourtiaceae (Chase et al2002 see also Judd 1997 Nandi et al 1998Savolainen et al 2000a) Salicaceae are considerablyexpanded to include flacourtiaceous taxa with salicoid

teeth (Nandi et al 1998) cocarcinogens and flowers inwhich the sepals and petals if both are present areequal in number However most of the taxa with cyclo-pentenoid cyanogenic glycosides and flowers in whichsepals and petals are not equal in number are placedin the newly accepted Achariaceae Sister to the rest ofSalicaceae is Casearia although this placement isonly weakly supported in Chase et al (2002 onlyrbcL) but strongly supported in a similar position withfar less taxonomic sampling but more data in Soltiset al (2000a three genes) Other families newlyrecognized here with genera that have been in Fla-courtiaceae sl include Lacistemataceae and Peridis-caceae Somewhat unexpectedly the poorly knownAchariaceae are sister to Kiggelaria (Soltis et al2000a Chase et al 2002) and so the name of the fam-ily becomes the conserved Achariaceae (not the olderbut non-conserved Kiggelariaceae as in several recentpapers) Other taxa with cyclopentenoid cyanogenicglycosides are Malesherbiaceae Turneraceae and Pas-sifloraceae The three are closely related (Chase et al2002) Turneraceae and Passifloraceae have foliarglands and biparental or paternal transmission ofplastids (eg Shore McQueen amp Little 1994) andMalesherbiaceae and Passifloraceae a corona Allthree possess a hypanthium-like structure that doesnot bear the stamens optional synonymization is thusappropriate

No molecular evidence supports Euphorbiaceae slas monophyletic and here they are divided into threefamilies (as in Chase et al 2002) Euphorbiaceae sscomprise the uniovulate Euphorbioideae Croto-noideae and Acalyphoideae Phyllanthaceae includethe biovulate Phyllanthoideae whereas Picroden-draceae include the biovulate Oldfieldioideae Thethree families have similar and distinctive fruits andsimilarities in embryology but other embryologicaldetails as well as features of leaf flower pollen andseed coat anatomy are distinct within each of the threefamilies

Linaceae are extended to include Hugoniaceae anda close relationship of the two has long been sug-gested Ochnaceae Medusagynaceae and Quiinaceaeform a distinctive and monophyletic group (Nandiet al 1998 Savolainen et al 2000a) with leaves hav-ing the secondary and tertiary venation particularlywell developed Optional synonymization seemsappropriate

Evidence provided by Litt amp Chase (1999 see alsoNandi et al 1998) strongly supports monophyly of agroup of four mostly small families TrigoniaceaeDichapetalaceae Chrysobalanaceae and Euphroni-aceae Optional recognition of an expanded Chrysobal-anaceae is recommended for these All havetenuinucellate ovules some species of each haveobliquely bisymmetric flowers and all have a single

408 AGP II


style The sister-group relationship of Erythroxylaceaeand Rhizophoraceae is confirmed by numerous fea-tures such as alkaloids and sieve tube plastid typethe rather poorly known African Aneulophus ofErythroxylaceae is particularly similar to some prim-itive Rhizophoraceae Optional synonymization isappropriate

In Oxalidales two alterations to APG (1998) havebeen made Brunelliaceae have been resurrected fromsynonymy because including them in Cunoniaceaewas premature Brunellia has been shown to berelated to both Cunoniaceae and Elaeocarpaceae(Bradford amp Barnes 2001 Savolainen et al 2000b)Tremandraceae (three genera from Australia) areembedded deeply in Elaeocarpaceae so the name ishere treated as a synonym of that family

In the eurosid II clade several minor changes havebeen made relative to APG (1998) Although Brassi-cales have remained unchanged here resurrection ofCapparaceae and Cleomaceae may be appropriate inthe future based on the results of Hall Sytsma amp Iltis(2002) who showed that Brassicaceae (sensu APG1998) comprise three strongly supported monophyl-etic groups representing Brassicaceae in the narrowsense Capparaceae subfamily Capparoideae and Cap-paraceae subfamily Cleomoideae They also point outthat there are some morphological features consistentwith this three-family view Emblingiaceae are placedin Brassicales based on the results of GregoryChandler amp Bayer (2000) We list Cochlospermaceaeas well as Diegodendronaceae as optional synonyms ofBixaceae Thymelaeaceae have likewise beenexpanded by the inclusion of Tepuianthus (Wurdack ampHorn 2001) the type of Tepuianthaceae which is wellsupported as sister to Thymelaeaceae Further work isneeded to evaluate relationships of Dipterocarpaceaeto Cistaceae and Sarcolaenaceae Dayanandan et al(1999) did not include Cistaceae and found an ambig-uous relationship of Dipterocarpaceae to Sarcolaen-aceae Savolainen et al (2000b) showed with rbcLdata that Pakaraimaea of Dipterocarpaceae isstrongly supported as sister to Cistus + Helianthe-mum and in all their shortest trees Monotes (Diptero-carpaceae the type of Monotaceae) was sister toSarcolaena (the type of Sarcolaenaceae) although thisreceived bootstrap support of less than 50 In Sap-indales Peganaceae are a possible synonym of Nitrar-iaceae both of which were at one time considered to bemembers of Zygophyllaceae (Sheahan amp Chase 19962000)

ASTERIDS

The asterids are a strongly supported monophyleticgroup including the same 10 orders as in APG (1998)Bremer et al (2002) analysed representatives of

amost all families of asterids using three genes (rbcLatpB and matK) and three non-coding plastid regionsand found Cornales to be the sister of all otherasterids followed by Ericales sister to the rest bothwith high jackknife percentages The rbcLatpB18SrDNA data (Soltis et al 2000a) indicated Cornales assister to Ericales whereas the ndhF data alone(Olmstead et al 2000) or ndhF together withrbcLatpB18S rDNA data (Albach et al 2001b)showed Cornales as sister to the rest of the asteridsbut without high support percentages Five families ofuncertain position in APG (1998) have been shown tobelong to the asterids Paracryphiaceae (of uncertainposition within the euasterid II clade as discussedunder Dipsacales) Pentaphylacaceae and Sladeni-aceae (the latter considered an optional synonym ofPentaphylacaceae of Ericales see below) Kaliphora-ceae (included in Montiniaceae of SolanalesSavolainen et al 2000a) and Cardiopteridaceae(Aquifoliales Karingrehed 2001) Furthermore recentanalyses support ordinal positions for several familiesthat were left unclassified to order in the APG systemalthough listed under euasterids I or II

Relationships within Cornales are still unclear butthe six families are all monophyletic In many studiesHydrostachys (formerly Hydrostachyaceae) has beenplaced within Hydrangeaceae (Soltis et al 2000aAlbach et al 2001a b) although the exact position ofthe genus within Hydrangeaceae is unclear In otherstudies it has fallen outside Hydrangeaceae (Xianget al 2002) It has been noted that for most genesHydrostachys has a great number of unique substitu-tions and the question of spurious attraction wasaddressed by Albach et al (2001a) Pending furtheranalyses we retain Hydrostachyaceae as a separatefamily Curtisia appears to be sister to Grubbiaceae(Soltis et al 2000a) not Cornaceae so Curtisiaceaeare here re-instated

Ericales comprise 23 families Relationships withinEricales have some structure but many relationshipsare still unclear One well-supported monophyleticgroup comprises Balsaminaceae Marcgraviaceae andTetrameristaceae (Soltis et al 2000a AnderbergRydin amp Kaumlllersjouml 2002 Bremer et al 2002 Tet-rameristaceae and the monogeneric Pellicieraceaehere being considered optional synonyms) it is sisterto the rest of the order Another well sup-portedgroup recently investigated in detail isthe primuloid group of families comprising thenewly re-circumscribed Primulaceae MyrsinaceaeTheophrastaceae and a new monogeneric familyMaesaceae (Anderberg Staringhl amp Kaumlllersjouml 2000Anderberg et al 2002 Kaumlllersjouml Bergqvist ampAnderberg 2000) A third group with robust support isformed by Actinidiaceae Roridulaceae Sarraceni-aceae Clethraceae Cyrillaceae and Ericaceae



(Anderberg et al 2002 Bremer et al 2002 classifica-tion of the last treated in Kron et al 2002) Styrac-aceae are sister to Diapensiaceae (94 jackknifesupport Anderberg et al 2002 Bremer et al 2002)and Halesia is nested within Styracaceae (Soltis et al2000a Anderberg et al 2002 Bremer et al 2002) soHalesiaceae are here reduced to synonymy under Sty-racaceae Pentaphylax appears as sister to Cardiopt-eris in the rbcL analysis of Savolainen et al 2000a)but analyses of sequences from five genes place Penta-phylax Ficalhoa and Sladenia with strong support inEricales (Anderberg et al 2002) The Savolainen et al(2000a) rbcL sequence for Pentaphylax was producedfrom highly degraded DNA extracted from herbariummaterial and seems to be a contaminant or an artifact(V Savolainen pers comm) Anderberg et al (2002)found that Sladenia and Ficalhoa are sister taxa (71jackknife support) and the two together are sister toTernstroemiaceae plus Pentaphylax (68 support)Ternstroemiaceae ss has 98 support and Pentaphy-lax together with Ternstroemiaceae ss has 97 sup-port (Anderberg et al 2002) Sladenia and Ficalhoawith their rather small flowers in cymose inflores-cences can be combined in Sladeniaceae (althoughFicalhoa has a straight embryo) but Anderberg et al(2002) proposed including them in Ternstroemiaceaealong with Pentaphylax which like other taxa of thatfamily has a curved embryo Lissocarpa (the type ofLissocarpaceae) is sister to Diospyros (100 support)and the two are usefully combined in an expandedEbenaceae Lissocarpa differing mainly in its inferiorovary (Berry et al 2001 Anderberg et al 2002Bremer et al 2002) Other less well supported groupsinclude Fouquieriaceae as sister to Polemoniaceae(72 in Anderberg et al 2002 88 in Bremer et al2002) and Sapotaceae as sister to Lecythidaceae sl(60 Anderberg et al 2002)

All euasterids are strongly supported as monophyl-etic and with the six DNA regions analysed byBremer et al (2002) euasterid I and II both receivedhigh jackknife percentages (100 and 99 respec-tively for which they also proposed the new informalnames of lamiids and campanulids) In earlier analy-ses both groups have low internal support EuasteridI had low jackknifebootstrap support 5366 (Olm-stead et al 2000) 56 (Soltis et al 2000a) or 40(Albach et al 2001b) and euasterid II has 68 (Olm-stead et al 2000) 88 (Soltis et al 2000a) or below33 (Albach et al 2001b) The percentages from thelatest study (Albach et al 2001b) are low and puz-zling because one would not expect lower scores ifdata sets are combined as was done in that study

In euasterid I there are some changes regardingfamilies not classified to order Recent investigationshave shown that Icacinaceae are polyphyletic(Savolainen et al 2000a Soltis et al 2000a Karingrehed

2001) and Doweld (2001) has recently segregatedmost of the genera as done here but assigned Emmo-tum to its own order and family Several genera infamilies listed in euasterid II by APG (1998) now showrelationships to Cardiopteridaceae and AquifolialesOther genera notably Icacina (Icacinaceae) belong toeuasterid I and have an apparent relationship(although not well supported) to Garryales PreviouslyAquifoliales included Oncothecaceae (APG 1998) butthat placement was premature as no internal supporthas been found for that position Icacinaceae andOncothecaceae are now listed under euasterid I butwithout an order as are Boraginaceae and VahliaceaeDespite several independent analyses based on multi-ple genes with broad taxon sampling there is so far noclear placement of Boraginaceae and Vahliaceae

Garryales now consist of Eucommiaceae and Garry-aceae Aucuba (the type of Aucubaceae) is hereincluded in Garryaceae In all molecular analysesGarrya and Aucuba have been sister taxa with highsupport and the molecular result is supported bymany morphological (pollen and embryological) andchemical characters (Bremer et al 2001) All mem-bers of Garryales have unisexual flowers and apicalplacentation which may be morphological synapomor-phies for the order

Gentianales still contain five families but circum-scription of some of these has been changed Logani-aceae were shown earlier to be polyphyletic anddetailed studies indicate that only 13 of the generabelong to the family (Backlund Oxelman amp Bremer2000) Other former Loganiaceae have been reas-signed to several other families The inclusion of Pte-leocarpa formerly Boraginaceae sl in Gelsemiaceaeis likely (Olmstead amp Ferguson 2001) Molecular datanow provide further support for inclusion of Dialypeta-lanthus (formerly Dialypetalanthaceae) within Rubi-aceae (Fay et al 2000a)

Lamiales are strongly supported as a monophyleticgroup of 23 families two of which were previously(APG 1998) not classified to order Plocospermata-ceae earlier unplaced in euasterid I are the sistergroup to the rest of Lamiales (Oxelman Backlund ampBremer 1999 Savolainen et al 2000a Bremer et al2002) and Carlemanniaceae have been shown to beclose to Oleaceae (Savolainen et al 2000a) Within theorder only a few interfamilial relationships receivedsupport including a few of the basal nodes Plocosper-mataceae are sister to the rest followed by Oleaceaetogether with Carlemanniaceae and subsequently Tet-rachondraceae as sister to the rest (Oxelman et al1999 Savolainen et al 2000a Olmstead et al 2000Bremer et al 2002) In spite of analyses involvingthree and more genes relationships among most fam-ilies remain unclear and more work needs to be doneIn APG (1998) Martyniaceae were listed as a syn-

410 AGP II


onym or sister taxon to Pedaliaceae but subsequentanalyses (Albach et al 2001b) have not found anysupport for this sister-group relationship and Mar-tynia is distant from Pedaliaceae in the trees Bremeret al (2002) found Avicennia to be nested in Acan-thaceae so Avicenniaceae is here included in Acan-thaceae A close relationship between Buddlejaceaeand Scrophulariaceae was suggested by Dahlgren(1983) based on chemical data but probably becauseof the artificial circumscription of both families involv-ing different unrelated entities they were kept sepa-rate In later analyses based on ndhF and rbcL data100 bootstrap support was found for a sister-grouprelationship between a restricted Buddlejaceae (Bud-dleja Emorya Gomphostigma and Nicodemia) andScrophulariaceae ss (Oxelman et al 1999) and thesame relationship was also supported when morpho-logical data were added (Bremer et al 2001) InOlmstead et al (2001 three genes) they also pre-sented support for a close relationship of these twofamilies with Myoporaceae and they included bothBuddlejaceae and Myoporaceae in Scrophulariaceae aclassification followed here The genus Androya ear-lier placed in Loganiaceae also belongs to the Myo-porum clade of the extended Scrophulariaceae Othergenera of the former Buddlejaceae andor Loganiaceaethat now belong to other families of Lamiales(Oxelman et al 1999) are Nuxia in Stilbaceae Peltan-thera and Sanango in Gesneriaceae and Polypre-mum in Tetrachondraceae A number of other generaremained unplaced to family but Mimulus appearscloser to Phryma than any genus now assigned toScrophulariaceae (Beardsley amp Olmstead 2002) sowe treat it there Parts of the former Scrophulariaceaehave also been transferred to Orobanchaceae andPlantaginaceae (Olmstead et al 2001) Cyclocheilonis nested in the expanded Orobanchaceae (Bremeret al 2002) so Cyclocheilaceae (= Nesogenaceae) arehere reduced to synonymy under Orobanchaceae

Solanales consist of five families of which three aresmall Of these Montiniaceae now including Kali-phora (the type of Kaliphoraceae Savolainen et al2000a) contain three small genera all characterizedby having unisexual flowers That character isunusual in euasterids but it occurs in a few genera indifferent families and is also common in Garryalesand Aquifoliales

In APG (1998) euasterid II included 10 families notclassified to order Two of these Icacinaceae and Car-lemanniaceae are now transferred to euasterid I andAdoxaceae are now included in Dipsacales (Bremeret al 2002) Parts of Icacinaceae remain among euas-terid II and the genera involved are now included inCardiopteridaceae and Stemonuraceae in Aquifoliales(Karingrehed 2001) There is no clear support for relation-ships among the families or between the unclassified

families and the orders but there is support for Ere-mosynaceae and Escalloniaceae as being closelyrelated (Hibsch-Jetter Soltis amp MacFarlane 1997Soltis et al 2000a Albach et al 2001a) The generaDesfontainia and Columellia are sister groups in Col-umelliaceae (optionally as two families APG 1998)In the analysis by Savolainen et al (2000a) the twogenera are unrelated The reasons for this are unclearand the sequences of Desfontainia and Columelliafrom GenBank fall together in other studies(Backlund et al 2000)

Aquifoliales are strongly supported as the sistergroup to the rest of euasterid II (Soltis et al 2000aBremer et al 2002) Cardiopteridaceae have beenexpanded to include several former genera of Icaci-naceae eg Gonocaryum Stemonuraceae haverecently been described and comprise a strongly sup-ported group of former genera of Icacinaceae eg Irv-ingbaileya (Karingrehed 2001)

Apiales have in recent investigations receivedstrong support as monophyletic (Olmstead et al 2000Soltis et al 2000a Bremer et al 2002) The ordernow comprises eight families with Pennantiaceaepreviously in Icacinaceae being included (Karingrehed2001 2003) The relationships among the small fami-lies of the order are still unclear and there are stilluncertainties about the delimitation of Apiaceae andAraliaceae (Plunkett amp Lowry 2001) Some of the fam-ilies are monogeneric and could possibly be mergedwhen well-supported sister-group relationships havebeen established Newly proposed Mackinlayaceaeand Myodocarpaceae include genera previously con-sidered to be archaic members of Araliaceae (seePlunkett 2001 Plunkett amp Lowry 2001 Karingrehed2003)

Asterales are strongly supported as monophyleticand contain 12 families Carpodetaceae are beingmerged with Rousseaceae (Lundberg 2001) Cyphiathe type of Cyphiaceae has appeared as sister to therest of Campanulaceae (optionally including Lobeli-aceae) in several recent rbcL analyses (eg Karingrehedet al 1999 Savolainen et al 2000a Lundberg 2001)However it appears that the rbcL sequence of Cyphiahitherto used is a pseudo-gene (Lundberg amp Bremer2002) and re-analysis with a new sequence placesCyphia as sister to other Lobeliaceae excluding Cam-panulaceae ss (see also Haberle 1998) Hence theoption of recognizing Campanulaceae and Lobeliaceaeas separate families is retained Interrelationshipsamong the families of Asterales are generally stilluncertain Since 1998 at least seven additional com-prehensive studies have included a wide family sam-pling of the asterids (Karingrehed et al 1999 Olmsteadet al 2000 Soltis et al 2000a Savolainen et al2000a Albach et al 2001b Bremer et al 2002Lundberg amp Bremer 2002) Unfortunately interrela-



tionships among families in Asterales in these studiesare somewhat different although in most cases thedifferences lack jackknifebootstrap support greaterthan 50 However Asteraceae Calyceraceae andGoodeniaceae together with their sister group Men-yanthaceae form a monophyletic group that isstrongly supported (Karingrehed et al 1999 Olmsteadet al 2000 Soltis et al 2000a Bremer et al 2002Lundberg amp Bremer 2002) The relationships amongthe first three families are unclear The rbcL and ndhFdata (Karingrehed et al 1999) and ndhF data (Olmsteadet al 2000) support Asteraceae and Calyceraceae assister families whereas rbcL together with atpB and18S rDNA (Soltis et al 2000a) support Goodeniaceaeand Calyceraceae as sister taxa With morphologicaldata rbcL ndhF and atpB sequences pooled there isstrong support for Asteraceae and Calyceraceae as sis-ter groups (Lundberg amp Bremer 2002) a result thatwas also obtained by Bremer et al (2002) in an anal-ysis of six DNA regions Another example of differentphylogenetic patterns of support between rbcLndhF(Karingrehed et al 1999) and rbcLatpB18S rDNA data(Soltis et al 2000a) is the well-supported relationshipbetween Argophyllaceae and Phellinaceae in the rbcLndhF analysis Stylidiaceae and Donatiaceae are close(Lundberg amp Bremer 2002) the latter is placed inoptional synonymy under the former

Dipsacales as here circumscribed are expanded toinclude Adoxaceae This family was unplaced in euas-terid II (APG 1998) but recent studies show supportfor an expanded circumscription (Soltis et al 2000aAlbach et al 2001b Bell et al 2001 Bremer et al2001 2002) In some recent systematics texts (egJudd et al 1999 2002) all other families of the orderwere merged into a single family Caprifoliaceae whichwe have indicated here as an option although somespecialists do not favour this broad concept All of thefamilies of Dipsacales originally in APG (1998) aremonophyletic none is monogeneric and some (egDipsacaceae and Valerianaceae) are well-knownentities with several hundred species Savolainen et al(2000a) included four additional families inDipsacales Desfontainiaceae (here included inColumelliaceae) Paracryphiaceae Polyosmaceae andSphenostemonaceae but there was no bootstrap sup-port for this expansion of Dipsacales so we retain thesefour families as unclassified to order Paracryphiaceaeare transferred to the euasterid II clade from the list offamilies of uncertain position (Bremer et al 2002)Both Paracryphiales and Desfontainiales are availableshould a name at an ordinal rank be required

CONCLUSION

We emphasize that the APG classification is proposedto facilitate communication we name organisms

because biologists require names for accurate commu-nication Progress since the first Angiosperm Phylog-eny Group consensus classification (APG 1998) hasbeen considerable Well-supported hypotheses of rela-tionships for many of the taxa that were unplacedthere have since been proposed and these ideas allowtheir assignment to orders of which five are newly rec-ognized here Furthermore the basic structure ofangiosperm phylogeny that was the foundation for theorders recognized in 1998 has been confirmed andstrengthened Nevertheless our knowledge of rela-tionships between many of the basal clades ofangiosperms among major eudicot lineages andmany orders such as Malpighiales and Lamialesremain to be resolved It is clear where we should con-centrate our efforts as only with a much more fullyresolved tree will we have a framework adequate tobegin to understand the details of morphological evo-lution of flowering plants Further progress in estab-lishing the relationships of clades will depend oncontinued broad collaboration

REFERENCES

Albach DC Soltis DE Chase MW Soltis PS 2001aPhylogenetic placement of the enigmatic angiospermHydrostachys Taxon 50 781ndash805

Albach DC Soltis PS Soltis DE Olmstead RG 2001bPhylogenetic analysis of the Asteridae based on sequences offour genes Annals of the Missouri Botanical Garden 88163ndash212

Anderberg AA Rydin C Kaumlllersjouml M 2002 Phylogeneticrelationships in the order Ericales s l analyses of moleculardata from five genes from the plastid and mitochondrialgenomes American Journal of Botany 89 677ndash687

Anderberg AA Staringhl B Kaumlllersjouml M 2000 Maesaceae anew primuloid family in the order Ericales sl Taxon 49183ndash197

APG 1998 An ordinal classification for the families of flower-ing plants Annals of the Missouri Botanical Garden 85 531ndash553

Appelquist WL Wallace RS 2000 Phylogeny of the Mada-gascan endemic family Didieraceae Plant Systematics andEvolution 221 157ndash166

Backlund M Oxelman B Bremer B 2000 Phylogeneticrelationships within the Gentianales based on ndhF andrbcL sequences with particular reference to the Logani-aceae American Journal of Botany 87 1029ndash1043

Barkman TJ Chenery G McNeal JR Lyons-Weiler JdePamphilis CW 2000 Independent and combined analy-ses of sequences from all three genomic compartments con-verge on the root of flowering plant phylogeny Proceedingsof the National Academy of Sciences USA 97 13166ndash13171

Beardsley PM Olmstead RG 2002 Redefining Phrymacaethe placement of Mimulus tribe Mimuleae and PhzrmaAmerican Journal of Botany 89 1093ndash1102

412 AGP II


Bell CD Edwards EJ Kim S-T Donoghue MJ 2001 Dip-sacales phylogeny based on chloroplast DNA sequencesHarvard Papers in Botany 6 481ndash489

Berry PE Savolainen V Sytsma KJ Hall JC Chase MW2001 Lissocarpa is sister to Diospyros (Ebenaceae) KewBulletin 56 725ndash729

Bortenschlager S 1973 Morphologie pollinique des Phyto-laccaceae Pollen et Spores 15 227ndash253

Bradford JC Barnes RW 2001 Phylogenetics and classifi-cation of Cunoniaceae (Oxalidales) using chloroplast DNAsequences and morphology Systematic Botany 26 354ndash385

Bremer K 2000 Phylogenetic nomenclature and the newordinal system of the angiosperms In Nordenstam B El-Ghazaly G Kassas M Laurent TC eds Plant systematics forthe 21st century London Portland Press 125ndash133

Bremer K 2002 Gondwanan evolution of the grass alliance offamilies (Poales) Evolution 56 1374ndash1387

Bremer K Backlund A Sennblad B Swenson UAndreasen K Hjertson M Lundberg J Backlund MBremer B 2001 A phylogenetic analysis of 100+ generaand 50+ families of euasterids based on morphological andmolecular data with notes on possible higher level morpho-logical synapomorphies Plant Systematics and Evolution229 137ndash169

Bremer B Bremer K Heidari N Erixon P AnderbergAA Olmstead RG Kaumlllersjouml M Barkhordarian E 2002Phylogenetics of asterids based on three coding and threenon-coding chloroplast DNA markers and the utility of non-coding DNA at higher taxonomic levels Molecular Phyloge-netics and Evolution 24 274ndash301

Briggs BG Johnson LAS 2000 Hopkinsiaceae and Lygini-aceae two new families of Poales in western Australia withrevisions of Hopkinsia and Lyginia Telopea 8 477ndash502

Caddick LR Rudall PJ Wilkin P Chase MW 2000 Yamsand their allies systematics of Dioscoreales In Wilson KLMorrison DA eds Systematics and evolution of monocotsProceedings of the 2nd International Monocot SymposiumMelbourne CSIRO 475ndash487

Caddick LR Rudall PJ Wilkin P Hedderson TAJ ChaseMW 2002a Phylogenetics of Dioscoreales based on com-bined analyses of morphological and molecular data Botan-ical Journal of the Linnean Society 138 123ndash144

Caddick LR Wilkin P Rudall PJ Hedderson TAJ ChaseMW 2002b Yams reclassified a recircumscription ofDioscoreaceae and Dioscoreales Taxon 51 103ndash114

Chase MW Duvall MR Hills HG Conran JG Cox AVEguiarte LE Hartwell J Fay MF Caddick LRCameron KM Hoot S 1995a Molecular systematics of Lil-ianae In Rudall PJ Cribb PJ Cutler DF Humphries CJeds Monocotyledons Systematics and Evolution Kew RoyalBotanic Gardens 109ndash137

Chase MW Soltis DE Olmstead RG Morgan D Les DHMishler BD Duvall MR Price RA Hills HG Qiu YLKron KA Rettig JH Conti E Palmer JD Manhart JRSytsma KJ Michael HJ Kress WJ Karol KA ClarkWD Hedreacuten M Gaut BS Jansen RK Kim KJ WimpeeCF Smith JF Furnier GR Strauss SH Xiang QY

Plunkett GM Soltis PS Swensen SM Williams SEGadek PA Quinn CJ Eguiarte LE Golenberg E LearnGH Graham SW Jr Barrett SCH Dayanandan SAlbert VA 1993 Phylogenetics of seed plants an analysis ofnucleotide sequences from the plastid gene rbcL Annals ofthe Missouri Botanical Garden 80 528ndash580

Chase MW Soltis DE Soltis PS Rudall PJ Fay MFHahn WJ Sullivan S Joseph J Molvray M Kores PJGivnish TJ Sytsma KJ Pires JC 2000 Higher-level sys-tematics of the monocotyledons An assessment of currentknowledge and a new classification In Wilson KL MorrisonDA eds Systematics and evolution of monocots Proceedingsof the 2nd International Monocot Symposium MelbourneCSIRO 3ndash16

Chase MW Stevenson WDW Wilkin P Rudall PJ 1995bMonocot systematics a combined analysis In Rudall PJCribb PJ Cutler DF Humphries CJ eds MonocotyledonsSystematics and evolution Kew Royal Botanic Gardens685ndash730

Chase MW Zmartzty S Lledoacute MD Wurdack KJSwensen SM Fay MF 2002 When in doubt put it in Fla-courtiaceae a molecular phylogenetic analysis based onplastid rbcL DNA sequences Kew Bulletin 57 141ndash181

Clausing G Renner SS 2001 Molecular phylogenetics ofMelastomataceae and Memecylaceae implications for char-acter evolution American Journal of Botany 88 486ndash498

Conti E Baum D Sytsma K 1999 Phylogeny of Cryptero-niaceae and related families implications for morphologyand biogeography In XVI International Botanical Congressabstracts St Louis Missouri Botanical Garden 250

Conti E Litt A Sytsma KJ 1996 Circumscription of Myr-tales and their relationships to other rosids evidence fromrbcL sequence data American Journal of Botany 83 221ndash233

Contreras VR Scogin R Philbrick CT 1993 A phytochem-ical study of selected Podostemaceae systematic implica-tions Aliso 13 513ndash520

Cronquist A 1981 An integrated system of classification offlowering plants New York Columbia University Press

Cueacutenoud P Savolainen V Powell M Grayer RJ ChaseMW 2002 Molecular phylogenetics of the Caryophyllalesbased on combined analyses of 18S rDNA and rbcL atpB andmatK sequences American Journal of Botany 89 132ndash144

Cusset C Cusset G 1988 Eacutetude sur les Podostemales 9Delimitation taxinomiques dans les Tristichaceae Bulletindu Museacuteum drsquoHistoire Naturelle Seacuteries 4 (10) 149ndash175

Dahlgren R 1983 General aspects of angiosperm evolutionand macrosystematics Nordic Journal of Botany 3 119ndash149

Dahlgren RMT Clifford HT Yeo PF 1985 The families ofthe monocotyledons structure evolution and taxonomyBerlin Spinger-Verlag

Dayanandan S Ashton PS Williams SM Primack RB1999 Phylogeny of the tropical tree family Dipterocarpaceaebased on nucleotide sequences of the chloroplast rbcL geneAmerican Journal of Botany 86 1182ndash1190

Doweld AB 2001 Tentamen Systematis Plantarum Vascular-ium (Tracheophytorum) Moscow GEOS

Doyle JA Endress PK 2000 Morphological phylogeneticanalysis of basal angiosperms comparison and combination



with molecular data International Journal of Plant Sciences161 (6 Suppl) S121ndashS153

Duvall MR Clegg MT Chase MW Clark WD Kress WJHills HG Eguiarte LE Smith JF Gaut BS Zimmer EALearn GH 1993 Phylogenetic hypotheses for the monocot-yledons constructed from rbcL sequences Annals of the Mis-souri Botanical Garden 80 607ndash619

Engler A 1930 Saxifragaceae In Engler A Prantl Keds Die natuumlrlichen Pflanzenfamilien 18a Leipzig WEngelman 74ndash226

Farris JS Albert VA Kaumlllersjouml M Lipscomb D Kluge AG1996 Parsimony jackknifing outperforms neighbor-joiningCladistics 12 99ndash124

Fay MF Bremer B Prance GT van der Bank M BridsonD Chase MW 2000a Plastid rbcL sequence data show Dia-lypetalanthus to be a member of Rubiaceae Kew Bulletin 55853ndash864

Fay MF Rudall PJ Sullivan S Stobart KL de BruijnAY Reeves G Qamaruz-Zaman F Hong W-PJoseph J Hahn WJ Conran JG Chase MW 2000bPhylogenetic studies of Asparagales based on four plasticDNA loci In Wilson KL Morrison DA eds Systematicsand evolution of monocots Proceedings of the 2nd Inter-national Monocot Symposium Melbourne CSIRO 360ndash371

Felsenstein J 1985 Confidence limits on phylogenies anapproach using the bootstrap Evolution 39 783ndash791

Fishbein M Hufford L Soltis DE 2003 Phylogeny of Sax-ifragales patterns of floral evolution and taxonomic revisionSystematic Botany in press

Fuse S Tamura MN 2000 A phylogenetic analysis of theplastid matK gene with emphasis on Melanthiaceae sensulato Plant Biology 2 415ndash427

Givnish TJ Evans TM Pires JC Sytsma KJ 1999Polyphyly and convergent morphological evolution inCommelinales and Commelinidae evidence from rbcLsequence data Molecular Phylogenetics and Evolution 12360ndash385

Graham SW Olmstead RG 2000 Evolutionary significanceof an unusual chloroplast DNA inversion found in two basalangiosperm lineages Current Genetics 37 183ndash188

Gregory T Chandler GT Bayer RJ 2000 Phylogeneticplacement of the enigmatic Western Australian genusEmblingia based on rbcL sequences Plant Species Biology15 67ndash72

Greuter W McNeill J Barrie FR Burdet HM DemoulinV Filgueiras TS Nicolson DH Silva PC Skog JETrehane P Turland NJ Hawksworth DL 2000 Inter-national code of botanical nomenclature (Saint Louis Code)adopted by the Sixteenth International Botanical CongressSt Louis Missouri July ndash August 1999 Regnum Vegetabile138 1ndash474

Haberle RC 1998 Phylogenetic systematics of Pseudonema-cladus and the North American cyphioids (Campanulaceaesensu lato) MSc Thesis Northern Arizona University

Hall JC Sytsma KJ Iltis HH 2002 Phylogeny of Cappar-aceae and Brassicaceae based on chloroplast sequence dataAmerican Journal of Botany 89 1826ndash1842

Hibsch-Jetter C Soltis DE MacFarlane TD 1997 Phylo-genetic analysis of Eremosyne pectinata (Saxifragaceae sl)based on rbcL sequence data Plant Systematics and Evolu-tion 204 225ndash232

Hillis DM 1996 Inferring complex phylogenies Nature 383130

Hoot SB Magallon-Puebla S Crane PR 1999 Phylogenyof basal eudicots based on three molecular data sets atpBrbcL and 18S nuclear ribosomal DNA sequences Annals ofthe Missouri Botanical Garden 86 119ndash131

Jaumlger-Zuumlrn I 1997 Embryological and floral studies in Wed-dellina squamulosa Tul (Podostemaceae Tristichoideae)Aquatic Botany 57 151ndash182

Jeong SC Ritchie NJ Myrold DD 1999 Molecular phylog-enies of plants and Frankia support multiple origins of act-inorhizal symbioses Molecular Phylogenetics and Evolution13 493ndash503

Judd WS 1997 The Flacourtiaceae in the southeasternUnited States Harvard Papers in Botany 1 65ndash79

Judd WS Campbell CS Kellogg EA Stevens PF 1999Plant systematics ndash a phylogenetic approach SunderlandMassachusetts Sinauer

Judd WS Campbell CS Kellogg EA Stevens PFDonoghue MJ 2002 Plant systematics ndash a phyloge-netic approach 2nd edn Sunderland MassachusettsSinauer

Kaumlllersjouml M Bergqvist G Anderberg A 2000 Genericrealignment in primuloid families of the Ericales s l a phy-logenetic analysis based on DNA sequences from three chlo-roplast genes and morphology American Journal of Botany87 1325ndash1341

Kaumlllersjouml M Farris JS Chase MW Bremer B Fay MFHumphries CJ Petersen G Seberg O Bremer K 1998Simultaneous parsimony jacknife analysis of 2538 rbcL DNAsequences reveals support for major clades of green plantsland plants seed plants and flowering plants Plant System-atics and Evolution 213 259ndash287

Karingrehed J 2001 Multiple origin of the tropical forest treefamily Icacinaceae American Journal of Botany 88 2259ndash2274

Karingrehed J 2003 The family Pennantiaceae and its relation-ships to Apiales Botanical Journal of the Linnean Society141 1ndash24

Karingrehed J Lundberg J Bremer B Bremer K 1999Evolution of the Australasian families AlseuosmiaceaeArgophyllaceae and Phellinaceae Systematic Botany 24660ndash682

Kron KA Judd WS Stevens PF Crayn DM AnderbergAA Gadek PA Quinn CJ Luteyn JL 2002 Phylogeneticclassification of Ericaceae molecular and morphological evi-dence Botanical Review 68 335ndash423

Litt A Chase MW 1999 The systematic position of Euphro-nia with comments on the position of Balanops an analysisbased on rbcL sequence data Systematic Botany 23 401ndash409

Lundberg J 2001 The asteralean affinity of the MauritianRoussea (Rousseaceae) Botanical Journal of the LinneanSociety 136 267ndash276

414 AGP II


Lundberg J Bremer K 2002 A phylogenetic study of theorder Asterales using one large morphological and threemolecular data sets International Journal of Plant Sciencesin press

Manhart JR Rettig JH 1994 Gene sequence data InBehnke H-D Mabry TJ eds Caryophyllales evolution andsystematics Berlin Springer Verlag 235ndash246

Meimberg H Dittrich P Bringmann G Schlauer JHeubl G 2000 Molecular phylogeny of Caryophyllidae slbased on matK sequences with special emphasis on carniv-orous taxa Plant Biology 2 218ndash228

Morgan DR Soltis DE 1993 Phylogenetic relationshipsamong members of the Saxifragaceae sensu lato based onrbcL sequence data Annals of the Missouri Botanical Gar-den 80 631ndash660

Nandi O Chase MW Endress PK 1998 A combined cladis-tic analysis of angiosperms using rbcL and non-moleculardata sets Annals of the Missouri Botanical Garden 85 137ndash212

Neyland R 2002 A phylogeny inferred from large subunit(26S) ribosomal DNA sequences suggests that Burmannialesis polyphyletic Australian Plant Research 15 19ndash28

Nickrent DL 2002 Oriacutegenes filogeneacuteticos de las plantasparaacutesitas In Loacutepez-Saacuteez JA Catalaacuten P Saacuteez L eds Plantasparaacutesitas de la Peniacutensula Ibeacuterica e Islas Baleares MadridSpain Mundi-Prensa Libros 29ndash56

Nickrent DL Blarer A Qiu Y-L Soltis DE Zanis M2001 Paleoherb status of Hydnoraceae supported by multi-gene analyses In Botany 2001 plants and peopleAbstracts Columbus Ohio Botanical Society of America130ndash131

Nickrent DL Duff RJ 1996 Molecular studies of parasiticplants using ribosomal RNA In Moreno MT Cubero JIBerner D Joel D Musselman LJ Parker C eds Advances inparasitic plant research Cordoba Spain Junta de Andalu-cia Direccion General de Investigacion Agraria 28ndash52

Nickrent DL Duff RJ Colwell AE Wolfe AD Young NDSteiner KE dePamphilis CW 1998 Molecular phyloge-netic and evolutionary studies of parasitic plants In SoltisDE Soltis PS Doyle JJ eds Molecular systematics of plantsII Boston Kluwer 211ndash241

Nickrent DL Maleacutecot V 2001 A molecular phylogeny ofSantalales In Fer A Thalouarn P Joel DM Musselman LJParker C Verkleij JAC eds Proceedings of the 7th Interna-tional Parasitic Weed Symposium Nantes France FaculteacuteDes Sciences Universiteacute de Nantes 69ndash74

Olmstead RG DePamphilis CW Wolfe AD Young NDElisons WJ Reeves PA 2001 Disintegration of theScrophulariaceae American Journal of Botany 88 348ndash361

Olmstead RG Ferguson D 2001 A molecular phylogeny ofthe Boraginaceae-Hydrophyllaceae In Botany 2001 plantsand people Abstracts Columbus Ohio Botanical Society ofAmerica 131

Olmstead RG Kim K-J Jansen RK Wagstaff SJ 2000The phylogeny of the Asteridae sensu lato based on chloro-plast ndhF gene sequences Molecular Phylogenetics andEvolution 16 96ndash112

Oxelman B Backlund M Bremer B 1999 Relationships ofBuddlejaceae sl investigated using parsimony jackknife andbranch support analysis of chloroplast ndhF and rbcLsequence data Systematic Botany 24 164ndash182

Pires JC Sytsma KJ 2002 A phylogenetic evaluation of abiosystematic framework Brodiaea and related petaloidmonocots (Themidaceae) American Journal of Botany 891342ndash1359

Plunkett GM 2001 Relationship of the order Apiales to sub-class Asteridae a re-evaluation of morphological charactersbased on insights from molecular data Edinburgh Journal ofBotany 8 183ndash200

Plunkett GM Lowry PP 2001 Relationships amonglsquoancient araliadsrsquo and their significance for the systematicsof Apiales Molecular Phylogenetics and Evolution 19 259ndash276

Qiu Y-L Chase MW Hoot SB Conti E Crane PR SytsmaKJ Parks CR 1998 Phylogenetics of Hamamelidae andtheir allies parsimony analyses of nucleotide sequences ofthe plastid gene rbcL International Journal of Plant Sci-ences 159 891ndash905

Qiu Y-L Lee J Bernasconi-Quadroni F Soltis DE SoltisPS Zanis M Zimmer EA Chen Z Savolainen V ChaseMW 1999 The earliest angiosperms evidence from mito-chondrial plastid and nuclear genomes Nature 402 404ndash407

Renner SS 1999 Circumscription and phylogeny of the Lau-rales evidence from molecular and morphological dataAmerican Journal of Botany 86 1301ndash1315

Reveal JL 1998ndashonward Indices nominum suprageneri-corum plantarum vascularium Alphabetical listing by gen-era of validly published suprageneric names httpwwwinformumdeduPBIOfaminspvindexhtml

Ronse Decraene LP Smets EF 1999 Similarities in floralontogeny and anatomy between the genera Francoa (Fran-coaceae) and Greyia (Greyiaceae) International Journal ofPlant Sciences 160 377ndash393

Rudall PJ Conran JG Chase MW 2000a Systematics ofRuscaceaeConvallariaceae a combined morphological andmolecular investigation Botanical Journal of the LinneanSociety 134 73ndash92

Rudall PJ Cribb PJ Cutler DF Humphries CJ 1995Monocotyledons systematics and evolution Kew RoyalBotanic Gardens

Rudall PJ Furness CA Fay MF Chase MW 2000b Con-sider the lilies ndash systematics of Liliales In Wilson KL Mor-rison DA eds Systematics and evolution of monocotsProceedings of the 2nd International Monocot SymposiumMelbourne CSIRO 347ndash359

Savolainen V Chase MW Hoot SB Morton CM SoltisDE Bayer C Fay MF de Bruijn AY Sullivan S QiuY-L 2000a Phylogenetics of flowering plants based oncombined analysis of plastid atpB and rbcL gene sequencesSystematic Biology 49 306ndash362

Savolainen V Fay MF Albach DC Backlund A van derBank M Cameron KM Johnson SA Lledoacute MDPintaud J-C Powell M Sheahan MC Soltis DE SoltisPS Weston P Whitten WM Wurdack KJ Chase MW2000b Phylogeny of the eudicots a nearly complete familial



analysis based on rbcL gene sequences Kew Bulletin 55257ndash309

Savolainen V Spichiger R Manen JF 1997 Polyphyletismof Celastrales deduced from a chloroplast non-coding DNAregion Molecular Phylogenetics and Evolution 7 145ndash157

Sheahan MC Chase MW 1996 A phylogenetic analysis ofZygophyllaceae R Br based on morphological anatomicaland rbcL sequence data Botanical Journal of the LinneanSociety 122 279ndash300

Sheahan MC Chase MW 2000 Phylogenetic relationshipswithin Zygophyllaceae based on DNA sequences of threeplastid regions with special emphasis on ZygophylloideaeSystematic Botany 25 371ndash384

Shore JS McQueen KL Little SL 1994 Inheritance ofplastid DNA in the Turnera ulmifolia complex AmericanJournal of Botany 81 1636ndash1639

Simmons MP Clevinger CC Savolainen V Archer RHMathews S Doyle JJ 2001 Phylogeny of the Celastraceaeinferred from phytochrome B gene sequence and morphol-ogy American Journal of Botany 88 313ndash325

Soltis DE Senters AE Kim S Thompson JD Soltis PSZanis MJ de Craene LS Endress PK Farris JS 2003Gunnerales are sister to other core eudicots and exhibit flo-ral features of early-diverging eudicots American Journal ofBotany 90 461ndash470

Soltis PS Soltis DE Chase MW 1999 Angiosperm phylog-eny inferred from multiple genes as a tool for comparativebiology Nature 402 402ndash404

Soltis DE Soltis PS Chase MW Mort ME Albach DCZanis M Savolainen V Hahn WH Hoot SB Fay MFAxtell M Swensen SM Prince LM Kress WJ NixonKC Farris JA 2000a Angiosperm phylogeny inferred from18S rDNA rbcL and atpB sequences Botanical Journal ofthe Linnean Society 133 381ndash461

Soltis DE Soltis PS Nickrent DL Johnson LA Hahn WJHoot SB Sweere JA Kuzoff RK Kron KA Chase MWSwensen SM Zimmer EA Chaw SM Gillespie LJKress WJ Sytsma KJ 1997 Angiosperm phylogenyinferred from 18S ribosomal DNA sequences Annals of theMissouri Botanical Garden 84 1ndash49

Soltis PS Soltis DE Zanis MJ Kim S 2000b Basal lin-eages of angiosperms Relationships and implications for flo-ral evolution International Journal of Plant Sciences 161 (6Suppl) S97ndashS107

Soltis DE Soltis PS 1997 Phylogenetic relationships inSaxifragaceae sensu lato a comparison of topologies basedon 18S rDNA and rbcL sequences American Journal ofBotany 84 504ndash522

Sosa V Chase MW 2003 Phylogenetics of Crossosomataceaebased on rbcL DNA sequence data Systematic Botany 27 inpress

Stevens PF 2001 Angiosperm phylogeny website httpwwwmobotorgMOBOTresearchAPweb

Sytsma KJ Morawetz J Pires JC Nepokroeff M ContiE Zjhra M Hall JC Chase MW 2002 Urticalean rosidscircumscription rosid ancestry and phylogenetics based onrbcL trnLF and ndhF sequences American Journal of Bot-any 89 1531ndash1546

Takhtajan AL 1997 Diversity and classification of floweringplants New York Columbia University Press

Thorne RF 1992 Classification and geography of the flower-ing plants Botanical Review 58 225ndash348

Thorne RF 2001 The classification and geography offlowering plants dicotyledons of the class Angiospermae(subclasses Magnoliidae Ranunculidae CaryophyllidaeDilleniidae Rosidae Asteridae and Lamiidae) BotanicalReview 66 441ndash647

Ueda K Kosuge H Tobe H 1997 A molecular phylogenyof Celtidaceae and Ulmaceae (Urticales) based on rbcLnucleotide sequences Journal of Plant Research 110 171ndash178

Wiegrefe SJ Sytsma KJ Guries RP 1998 The Ulmaceaeone family or two Evidence from chloroplast DNA restric-tion site mapping Plant Systematics and Evolution 210249ndash270

Wilson KL Morrison DA 2000 Systematics and evolution ofmonocots Proceedings of the 2nd International MonocotSymposium Melbourne CSIRO

Wu C-Y Tang Y-C Chen Z-D Li D-Z 2002 Synopsis of anew lsquopolyphyletic-polychronic-polytopicrsquo system of theangiosperms Acata Phytotaxonoica Sinica 40 289ndash322

Wurdack KJ Horn JW 2001 A reevaluation of the affinitiesof the Tepuianthaceae molecular and morphological evi-dence for placement in the Malvales In Botany 2001 plantsand people Abstracts Columbus Ohio Botanical Society ofAmerica 151

Xiang Q-Y Moody M Soltis DE Fan CZ Soltis PS 2002Relationships within Cornales and circumscription of Cor-naceae ndash matK and rbcL sequence data and effects of out-groups and long branches Molecular Phylogenetics andEvolution 24 35ndash47

Zanis MJ Soltis DE Soltis PS Qiu Y-L Mathews SDonoghue MJ 2002 The root of the angiosperms revisitedProceedings of the National Academy of Sciences (USA) 996848ndash6853

Zanis MJ Soltis DE Soltis PS Qiu Y-L Zimmer EA 2003Phylogenetic analyses and perianth evolution in basalangiosperms Annals of the Missouri Botanical Garden inpress

APPENDIX

CLASSIFICATION OF FLOWERING PLANTS

The state of family name and authorships currently isin flux The International Code of Botanical Nomen-clature (Greuter et al 2000) provides currently for theuse of pre-1789 names However there is a majorpush which in all likelihood will be successful toestablish a formal starting date for spermatophyte (ifnot all vascular plants) family names as of 4 August1789 (eg Jussieursquos Genera plantarum) As a resultthis listing in an effort to avoid the introduction ofnames andor authorships that almost certainly willbe incorrect after 2005 presumes 1789 as the startdate for angiosperm family names In this way we

416 AGP II


believe nomenclatural stability can be achieved with-out undue confusion in the future Two names areretained (Potamogetonaceae and Cornaceae) in antic-ipation of future superconservation proposals formallyestablishing their continued use Also Meerow andothers likely will make a similar proposal to maintainAmaryllidaceae over Alliaceae but Alliaceae isretained here

new family placement daggernewly recognized orderfor the APG system sectnew family circumscriptiondescribed in the text The list reflects a starting datefor all flowering plant family names of 4 August 1789(Jussieu Genera plantarum) Full citations are avail-able elsewhere (Reveal 1998-onward) Families insquare brackets are acceptable monophyletic alterna-tives to the broader circumscription favoured here

Amborellaceae Pichon (1948) nom consChloranthaceae RBr ex Sims (1820) nom consNymphaeaceae Salisb (1805) nom cons[+Cabombaceae Rich ex ARich (1822) nom

cons]

daggerAustrobaileyales Takht ex Reveal (1992)Austrobaileyaceae (Croizat) Croizat (1943) nom

conssectSchisandraceae Blume (1830) nom cons[+Illiciaceae ACSm (1947) nom cons]Trimeniaceae LSGibbs (1917) nom cons

Ceratophyllales Bisch (1839)Ceratophyllaceae Gray (1821) nom cons

MAGNOLIIDS

daggerCanellales Cronquist (1957)Canellaceae Mart (1832) nom consWinteraceae RBr ex Lindl (1830) nom cons

Laurales Perleb (1826)Atherospermataceae RBr (1814)Calycanthaceae Lindl (1819) nom consGomortegaceae Reiche (1896) nom consHernandiaceae Bercht amp JPresl (1820) nom

consLauraceae Juss (1789) nom consMonimiaceae Juss (1809) nom consSiparunaceae (ADC) Schodde 1970

Magnoliales Bromhead (1838)Annonaceae Juss (1789) nom consDegeneriaceae IWBailey amp ACSm (1942) nom

consEupomatiaceae Endl (1841) nom consHimantandraceae Diels (1917) nom consMagnoliaceae Juss (1789) nom cons

Myristicaceae RBr (1810) nom cons

Piperales Dumort (1829)Aristolochiaceae Juss (1789) nom consHydnoraceae CAgardh (1821) nom consLactoridaceae Engl (1888) nom consPiperaceae Bercht amp J Presl (1820) nom consSaururaceae Martynov (1820) nom cons

MONOCOTS

sectPetrosaviaceae Hutch (1934) nom cons

Acorales Reveal (1996)Acoraceae Martynov (1820)

Alismatales Dumort (1829)Alismataceae Vent (1799) nom consAponogetonaceae JAgardh (1858) nom consAraceae Juss (1789) nom consButomaceae Mirb (1804) nom consCymodoceaceae NTaylor (1909) nom consHydrocharitaceae Juss (1789) nom consJuncaginaceae Rich (1808) nom consLimnocharitaceae Takht ex Cronquist (1981)Posidoniaceae Hutch (1934) nom consPotamogetonaceae Rchb (1828) nom consRuppiaceae Horan (1834) nom consScheuchzeriaceae FRudolphi (1830) nom consTofieldiaceae Takht (1995)Zosteraceae Dumort (1829) nom cons

Asparagales Bromhead (1838)sectAlliaceae Batsch ex Borkh (1797) nom cons[+Agapanthaceae FVoigt (1850)][+Amaryllidaceae JSt-Hil (1805) nom cons]sectAsparagaceae Juss (1789) nom cons[+Agavaceae Dumort (1829) nom cons][+Aphyllanthaceae Burnett (1835)][+Hesperocallidaceae Traub (1972)][+Hyacinthaceae Batsch ex Borkh (1797)][+Laxmanniaceae Bubani (1901 - 02)][+Ruscaceae Spreng (1826) nom cons][+Themidaceae Salisb (1866)]Asteliaceae Dumort (1829)Blandfordiaceae RDahlgren amp Clifford (1985)Boryaceae (Baker) MWChase Rudall amp Conran

(1997)Doryanthaceae RDahlgren amp Clifford (1985)Hypoxidaceae RBr (1814) nom consIridaceae Juss (1789) nom consIxioliriaceae Nakai (1943)Lanariaceae HHuber ex RDahlgren amp AEvan

Wyk (1988)Orchidaceae Juss (1789) nom consTecophilaeaceae Leyb (1862) nom cons



sectXanthorrhoeaceae Dumort (1829) nom cons[+Asphodelaceae Juss (1789)][+Hemerocallidaceae RBr (1810)]Xeronemataceae MWChase Rudall amp MFFay

(2001)

Dioscoreales Hookf (1873)sectBurmanniaceae Blume (1827) nom conssectDioscoreaceae RBr (1810) nom consNartheciaceae Fr ex Bjurzon (1846)

Liliales Perleb (1826)Alstroemeriaceae Dumort (1829) nom consCampynemataceae Dumort (1829)Colchicaceae DC (1804) nom consCorsiaceae Becc (1878) nom consLiliaceae Juss (1789) nom consLuzuriagaceae Lotsy (1911)Melanthiaceae Batsch ex Borkh (1796) nom

consPhilesiaceae Dumort (1829) nom consRhipogonaceae Conran amp Clifford (1985)Smilacaceae Vent (1799) nom cons

Pandanales Lindl (1833)Cyclanthaceae Poit ex ARich (1824) nom consPandanaceae RBr (1810) nom consStemonaceae Caruel (1878) nom consTriuridaceae Gardner (1843) nom consVelloziaceae Hook (1827) nom cons

COMMELINIDS

Dasypogonaceae Dumort (1829)

Arecales Bromhead (1840)Arecaceae Schultz Sch (1832) nom cons

Commelinales Dumort (1829)Commelinaceae Mirb (1804) nom consHaemodoraceae RBr (1810) nom consHanguanaceae Airy Shaw (1964)Philydraceae Link (1821) nom consPontederiaceae Kunth (1816) nom cons

Poales Small (1903)Anarthriaceae DFCutler amp Airy Shaw (1965)Bromeliaceae Juss (1789) nom consCentrolepidaceae Endl (1836) nom consCyperaceae Juss (1789) nom consEcdeiocoleaceae DFCutler amp Airy Shaw (1965)Eriocaulaceae Martynov (1820) nom consFlagellariaceae Dumort (1829) nom consHydatellaceae UHamann (1976)Joinvilleaceae Toml amp ACSm (1970)Juncaceae Juss (1789) nom cons

Mayacaceae Kunth (1842) nom consPoaceae (RBr) Barnh 1895 nom consRapateaceae Dumort (1829) nom consRestionaceae RBr (1810) nom consSparganiaceae Hanin (1811) nom conssectThurniaceae Engl (1907) nom consTyphaceae Juss (1789) nom conssectXyridaceae CAgardh (1823) nom cons

Zingiberales Griseb (1854)Cannaceae Juss (1789) nom consCostaceae Nakai (1941)Heliconiaceae Nakai (1941)Lowiaceae Ridl (1924) nom consMarantaceae RBr (1814) nom consMusaceae Juss (1789) nom consStrelitziaceae Hutch (1934) nom consZingiberaceae Martynov (1820) nom cons

EUDICOTS

sectBuxaceae Dumort (1822) nom cons[+Didymelaceae Leandri (1937)]Sabiaceae Blume (1851) nom consTrochodendraceae Eichler (1865) nom cons[+Tetracentraceae ACSm (1945) nom cons]

Proteales Dumort (1829)Nelumbonaceae Bercht amp JPresl (1820) nom

conssectProteaceae Juss (1789) nom cons[+Platanaceae TLestib (1826) nom cons]

Ranunculales Dumort (1829)Berberidaceae Juss (1789) nom consCircaeasteraceae Hutch (1926) nom cons[+Kingdoniaceae ASFoster ex Airy Shaw (1964)]Eupteleaceae KWilh (1910) nom consLardizabalaceae RBr (1821) nom consMenispermaceae Juss (1789) nom consPapaveraceae Juss (1789) nom cons[+Fumariaceae Bercht amp JPresl (1820) nom

cons][+Pteridophyllaceae (Murb) Nakai ex Reveal amp

Hoogland (1991)]Ranunculaceae Juss (1789) nom cons

CORE EUDICOTS

Aextoxicaceae Engl amp Gilg (1920) nom consBerberidopsidaceae Takht (1985)Dilleniaceae Salisb (1807) nom cons

daggerGunnerales Takht ex Reveal (1992)sectGunneraceae Meisn (1842) nom cons[+Myrothamnaceae Nied (1891) nom cons]

418 AGP II


Caryophyllales Perleb (1826)Achatocarpaceae Heimerl (1934) nom consAizoaceae Martynov (1820) nom consAmaranthaceae Juss (1789) nom consAncistrocladaceae Planch ex Walp (1851) nom

consAsteropeiaceae (Szyszyl) Takht ex Reveal amp

Hoogland (1990)Barbeuiaceae Nakai (1942)Basellaceae Raf (1837) nom consCactaceae Juss (1789) nom consCaryophyllaceae Juss (1789) nom consDidiereaceae Radlk (1896) nom consDioncophyllaceae Airy Shaw (1952) nom consDroseraceae Salisb (1808) nom consDrosophyllaceae Chrtek Slaviacutekovaacute amp Studnicka

(1989)Frankeniaceae Desv (1817) nom consGisekiaceae Nakai (1942)Halophytaceae ASoriano (1984)Molluginaceae Bartl (1825) nom consNepenthaceae Bercht amp JPresl (1820) nom

consNyctaginaceae Juss (1789) nom consPhysenaceae Takht (1985)Phytolaccaceae RBr (1818) nom consPlumbaginaceae Juss (1789) nom consPolygonaceae Juss (1789) nom consPortulacaceae Juss (1789) nom consRhabdodendraceae Prance (1968)Sarcobataceae Behnke (1997)Simmondsiaceae Tiegh (1899)Stegnospermataceae Nakai (1942)Tamaricaceae Bercht amp JPresl (1820) nom cons

Santalales Dumort (1829)Olacaceae RBr (1818) nom consOpiliaceae Valeton (1886) nom consLoranthaceae Juss (1808) nom consMisodendraceae J Agardh (1858) nom consSantalaceae RBr (1810) nom cons

Saxifragales Dumort (1829)Altingiaceae Horan (1843) nom consAphanopetalaceae Doweld (2001)Cercidiphyllaceae Engl (1907) nom consCrassulaceae JSt-Hil (1805) nom consDaphniphyllaceae Muumlll-Arg (1869) nom consGrossulariaceae DC (1805) nom conssectHaloragaceae RBr (1814) nom cons[+Penthoraceae Rydb ex Britt (1901) nom cons][+Tetracarpaeaceae Nakai (1943)]Hamamelidaceae RBr (1818) nom conssectIteaceae JAgardh (1858) nom cons[+Pterostemonaceae Small (1905) nom cons]Paeoniaceae Raf (1815) nom consSaxifragaceae Juss (1789) nom cons

ROSIDS

Aphloiaceae Takht (1985)Geissolomataceae Endl (1841)Ixerbaceae Griseb (1854)Picramniaceae Fernando amp Quinn (1995)Strasburgeriaceae Soler (1908) nom consVitaceae Juss (1789) nom cons

daggerCrossosomatales Takht ex Reveal (1993)Crossosomataceae Engl (1897) nom consStachyuraceae JAgardh (1858) nom consStaphyleaceae Martynov (1820) nom cons

Geraniales Dumort (1829)Geraniaceae Juss (1789) nom cons[+Hypseocharitaceae Wedd (1861)]Ledocarpaceae Meyen (1834)sectMelianthaceae Bercht amp JPresl (1820) nom

cons[+Francoaceae AJuss (1832) nom cons]Vivianiaceae Klotzsch (1836)

Myrtales Rchb (1828)Alzateaceae SAGraham (1985)Combretaceae RBr (1810) nom consCrypteroniaceae ADC (1868) nom consHeteropyxidaceae Engl amp Gilg (1920) nom consLythraceae JSt-Hil (1805) nom conssectMelastomataceae Juss (1789) nom cons[+Memecylaceae DC (1827) nom cons]Myrtaceae Juss (1789) nom consOliniaceae Arn (1839) nom consOnagraceae Juss (1789) nom consPenaeaceae Sweet ex Guill (1828) nom consPsiloxylaceae Croizat (1960)Rhynchocalycaceae LASJohnson amp BGBriggs

(1985)Vochysiaceae ASt-Hil (1820) nom cons

EUROSIDS I

sectZygophyllaceae RBr (1814) nom cons[+Krameriaceae Dumort (1829) nom cons]Huaceae AChev (1947)

daggerCelastrales Baskerville (1839)sectCelastraceae RBr (1814) nom consdaggerLepidobotryaceae JLeacuteonard (1950) nom consParnassiaceae Martynov (1820) nom cons[+Lepuropetalaceae Nakai (1943)]

Cucurbitales Dumort (1829)Anisophylleaceae Ridl (1922)Begoniaceae Bercht amp JPresl (1820) nom consCoriariaceae DC (1824) nom cons



Corynocarpaceae Engl (1897) nom consCucurbitaceae Juss (1789) nom consDatiscaceae Bercht amp JPresl (1820) nom consTetramelaceae Airy Shaw (1964)

Fabales Bromhead (1838)Fabaceae Lindl (1836) nom consPolygalaceae Hoffmanns amp Link (1809) nom

consQuillajaceae DDon (1831)Surianaceae Arn (1834) nom cons

Fagales Engl (1892)Betulaceae Gray (1821) nom consCasuarinaceae RBr (1814) nom consFagaceae Dumort (1829) nom conssectJuglandaceae DC ex Perleb (1818) nom cons[+Rhoipteleaceae Hand-Mazz (1932) nom cons]Myricaceae ARich ex Kunth (1817) nom consNothofagaceae Kuprian (1962)Ticodendraceae Goacutemez-Laur amp LDGoacutemez (1991)

Malpighiales Mart (1835)sectAchariaceae Harms (1897) nom consBalanopaceae Benth amp Hookf (1880) nom consBonnetiaceae (Bartl) LBeauv ex Nakai (1948)Caryocaraceae Voigt (1845) nom conssectChrysobalanaceae RBr (1818) nom cons[+Dichapetalaceae Baill (1886) nom cons][+Euphroniaceae Marc-Berti (1989)][+Trigoniaceae Endl (1841) nom cons]sectClusiaceae Lindl (1836) nom consCtenolophonaceae (HWinkl) Exell amp Mendonccedila

(1951)Elatinaceae Dumort (1829) nom conssectEuphorbiaceae Juss (1789) nom consGoupiaceae Miers (1862)Humiriaceae AJuss (1829) nom conssectHypericaceae Juss (1789) nom consIrvingiaceae (Engl) Exell amp Mendonccedila (1951)

nom consIxonanthaceae Planch ex Miq (1858) nom consLacistemataceae Mart (1826) nom conssectLinaceae DC ex Perleb (1818) nom consLophopyxidaceae (Engl) HPfeiff (1951)Malpighiaceae Juss (1789) nom conssectOchnaceae DC (1811) nom cons[+Medusagynaceae Engl amp Gilg (1924) nom

cons][+Quiinaceae Choisy ex Engl (1888) nom cons]Pandaceae Engl amp Gilg (1912ndash13) nom conssectPassifloraceae Juss ex Roussel (1806) nom

cons[+Malesherbiaceae DDon (1827) nom cons][+Turneraceae Kunth ex DC (1828) nom cons]Peridiscaceae Kuhlm (1950) nom cons

sectPhyllanthaceae Martynov (1820)sectPicrodendraceae Small (1917) nom consPodostemaceae Rich ex C Agardh (1822) nom

consPutranjivaceae Endl (1841)sectRhizophoraceae Pers (1807) nom cons[+Erythroxylaceae Kunth (1822) nom cons]sectSalicaceae Mirb (1815) nom consViolaceae Batsch (1802) nom cons

Oxalidales Heintze (1927)sectBrunelliaceae Engl (1897) nom consCephalotaceae Dumort (1829) nom consConnaraceae RBr (1818) nom consCunoniaceae RBr (1814) nom conssectElaeocarpaceae Juss ex DC (1816) nom consOxalidaceae RBr (1818) nom cons

Rosales Perleb (1826)Barbeyaceae Rendle (1916) nom conssectCannabaceae Martynov (1820) nom consDirachmaceae Hutch (1959)Elaeagnaceae Juss (1789) nom consMoraceae Link (1831) nom consRhamnaceae Juss (1789) nom consRosaceae Juss (1789) nom consUlmaceae Mirb (1815) nom conssectUrticaceae Juss (1789) nom cons

EUROSIDS II

Tapisciaceae (Pax) Takht (1987)

Brassicales Bromhead (1838)Akaniaceae Stapf (1912) nom cons[+Bretschneideraceae Engl amp Gilg (1924) nom

cons]Bataceae Perleb (1838) nom consBrassicaceae Burnett (1835) nom consCaricaceae Dumort (1829) nom consEmblingiaceae Airy Shaw (1964)Gyrostemonaceae Endl (1841) nom consKoeberliniaceae Engl (1895) nom consLimnanthaceae RBr (1833) nom consMoringaceae Martynov (1820) nom consPentadiplandraceae Hutch amp Dalziel (1928)Resedaceae Bercht amp JPresl (1820) nom consSalvadoraceae Lindl (1836) nom consSetchellanthaceae Iltis (1999)Tovariaceae Pax (1891) nom consTropaeolaceae Bercht amp JPresl (1820) nom

cons

Malvales Dumort (1829)sectBixaceae Kunth (1822) nom cons[+Diegodendraceae Capuron (1964)]

420 AGP II


[+Cochlospermaceae Planch (1847) nom cons]Cistaceae Juss (1789) nom consDipterocarpaceae Blume (1825) nom consMalvaceae Juss (1789) nom consMuntingiaceae CBayer MWChase amp MFFay

(1998)Neuradaceae Link (1831) nom consSarcolaenaceae Caruel (1881) nom consSphaerosepalaceae (Warb) Tiegh ex Bullock

(1959)sectThymelaeaceae Juss (1789) nom cons

Sapindales Dumort (1829)Anacardiaceae RBr (1818) nom consBiebersteiniaceae Endl (1841)Burseraceae Kunth (1824) nom consKirkiaceae (Engl) Takht (1967)Meliaceae Juss (1789) nom conssectNitrariaceae Bercht amp JPresl (1820) nom

cons[+Peganaceae (Engl) Tieghm ex Takht (1987)][+Tetradiclidaceae (Engl) Takht 1986)Rutaceae Juss (1789) nom consSapindaceae Juss (1789) nom consSimaroubaceae DC (1811) nom cons

ASTERIDS

Cornales Dumort (1829)Cornaceae Dumort (1829) nom cons[+Nyssaceae Juss ex Dumort (1829) nom cons]Curtisiaceae (Engl) Takht (1987)Grubbiaceae Endl (1839) nom consHydrangeaceae Dumort (1829) nom consHydrostachyaceae (Tul) Engl (1894) nom consLoasaceae Juss (1804) nom cons

Ericales Dumort (1829)Actinidiaceae Gilg amp Werderm (1825) nom consBalsaminaceae Bercht amp JPresl (1820) nom

consClethraceae Klotzsch (1851) nom consCyrillaceae Endl (1841) nom consDiapensiaceae Lindl (1836) nom conssectEbenaceae Guumlrke (1891) nom consEricaceae Juss (1789) nom consFouquieriaceae DC (1828) nom consLecythidaceae ARich (1825) nom consMaesaceae (ADC) Anderb BStaringhl amp Kaumlllersjouml

(2000)Marcgraviaceae Juss ex DC (1816) nom conssect Myrsinaceae RBr (1810) nom consPentaphylacaceae Engl (1897) nom cons[+Ternstroemiaceae Mirb ex DC (1816)][+Sladeniaceae Airy Shaw (1964)]Polemoniaceae Juss (1789) nom cons

sectPrimulaceae Batsch ex Borkh (1797) nomcons

Roridulaceae Bercht amp JPresl (1820) nom consSapotaceae Juss (1789) nom consSarraceniaceae Dumort (1829) nom conssectStyracaceae DC amp Spreng (1821) nom consSymplocaceae Desf (1820) nom conssectTetrameristaceae Hutch (1959)[+Pellicieraceae (Triana amp Planch) LBeauvis ex

Bullock (1959)]Theaceae Mirb ex Ker Gawl (1816) nom conssectTheophrastaceae Link (1829) nom cons

EUASTERIDS I

Boraginaceae Juss (1789) nom conssectIcacinaceae (Benth) Miers (1851) nom consOncothecaceae Kobuski ex Airy Shaw (1964)Vahliaceae Dandy (1959)

Garryales Lindl (1846)Eucommiaceae Engl (1909) nom conssectGarryaceae Lindl (1834) nom cons[+Aucubaceae JAgardh (1858)]

Gentianales Lindl (1833)Apocynaceae Juss (1789) nom consGelsemiaceae (GDon) Struwe amp V Albert (1995)Gentianaceae Juss (1789) nom consLoganiaceae RBr (1814) nom consRubiaceae Juss (1789) nom cons

Lamiales Bromhead (1838)sectAcanthaceae Juss (1789) nom consBignoniaceae Juss (1789) nom consByblidaceae (Engl amp Gilg) Domin (1922) nom

consCalceolariaceae (DDon) Olmstead (2001)Carlemanniaceae Airy Shaw (1964)Gesneriaceae Rich amp Juss ex DC (1816) nom

consLamiaceae Martynov (1820) nom consLentibulariaceae Rich (1808) nom consMartyniaceae Horan (1847) nom consOleaceae Hoffmanns amp Link (1809) nom consSee Orobanchaceae Vent (1799) nom consPaulowniaceae Nakai (1949)Pedaliaceae RBr (1810) nom conssectPhrymaceae Schauer (1847) nom conssectPlantaginaceae Juss (1789) nom consPlocospermataceae Hutch (1973)Schlegeliaceae (AHGentry) Reveal (1996)sectScrophulariaceae Juss (1789) nom consStilbaceae Kunth (1831) nom consTetrachondraceae Wettst (1924)Verbenaceae JSt-Hil (1805) nom cons



Solanales Dumort (1829)Convolvulaceae Juss (1789) nom consHydroleaceae Bercht amp J Presl (1820)sectMontiniaceae Nakai (1943) nom consSolanaceae Juss (1789) nom consSphenocleaceae (Lindl) Baskerville (1839) nom

cons

EUASTERIDS II

Bruniaceae Bercht amp JPresl (1820) nom consColumelliaceae DDon (1828) nom cons[+Desfontainiaceae Endl (1841) nom cons]Eremosynaceae Dandy (1959)Escalloniaceae RBr ex Dumort (1829) nom

consParacryphiaceae Airy Shaw (1964)Polyosmaceae Blume (1851)Sphenostemonaceae PRoyen amp Airy Shaw (1972)Tribelaceae Airy Shaw (1964)

Apiales Nakai (1930)Apiaceae Lindl (1836) nom consAraliaceae Juss (1789) nom consAralidiaceae Philipson amp BCStone (1980)Griseliniaceae JRForst amp GForst ex ACunn

(1839)Mackinlayaceae Doweld (2001)Melanophyllaceae Takht ex Airy Shaw (1972)Myodocarpaceae Doweld (2001)Pennantiaceae JAgardh (1858)Pittosporaceae RBr (1814) nom consTorricelliaceae Hu (1934)

Aquifoliales Senft (1856)Aquifoliaceae DC ex ARich (1828) nom conssectCardiopteridaceae Blume (1847) nom consHelwingiaceae Decne (1836)Phyllonomaceae Small (1905)sectStemonuraceae (M Roem) Karingrehed (2001)

Asterales Lindl (1833)Alseuosmiaceae Airy Shaw (1964)Argophyllaceae (Engl) Takht 1987Asteraceae Martynov (1820) nom consCalyceraceae RBr ex Rich (1820) nom conssectCampanulaceae Juss (1789) nom cons[+Lobeliaceae Juss ex Bonpl (1813) nom cons]Goodeniaceae RBr (1810) nom consMenyanthaceae Bercht amp JPresl (1820) nom

consPentaphragmataceae JAgardh (1858) nom consPhellinaceae (Loes) Takht 1967sectRousseaceae DC (1839)Stylidiaceae RBr (1810) nom cons[+Donatiaceae BChandler (1911) nom cons]

Dipsacales Dumort (1829)Adoxaceae EMey (1839) nom conssectCaprifoliaceae Juss (1789) nom cons[+Diervillaceae (Raf) Pyck 1998)[+Dipsacaceae Juss (1789) nom cons][+Linnaeaceae (Raf) Backlund 1998)[+Morinaceae Raf (1820)][+Valerianaceae Batsch (1802) nom cons]

TAXA OF UNCERTAIN POSITIONIf an unplaced genus is the type of a family name thatname is given for information purposes

Aneulophus BenthApodanthaceae van Tieghem ex Takhtajan in

Takhtajan (1997) [three genera]Bdallophyton EichlBalanophoraceae Rich (1822) nom consCentroplacus PierreCynomorium L [Cynomoriaceae Lindl (1833)

nom cons]Cytinus L [Cytinaceae ARich (1824)]Dipentodon Dunn [Dipentodontaceae Merr

(1941) nom cons]Gumillea Ruiz amp PavHoplestigma Pierre [Hoplestigmataceae Engl amp

Gilg (1924) nom cons]Leptaulus BenthMedusandra Brenan [Medusandraceae Brenan

(1952) nom cons]Metteniusa HKarst [Metteniusaceae HKarst

ex Schnizl (1860ndash1870)]Mitrastema Makino [Mitrastemonaceae Makino

(1911) nom cons]Pottingeria Prain [Pottingeriaceae (Engl) Takht

1987)Rafflesiaceae Dumort (1829) nom cons [three

genera included]Soyauxia OlivTrichostephanus Gilg

ORDINAL NAMES AND SYNONYMSAccepted ordinal names are in bold face those basedon a family not yet placed in an order are in italicsYear of publication is indicated

Acanthales Lindl (1833) = LamialesAcerales Lindl (1833) = SapindalesAcorales Reveal (1996)Actinidiales Takht ex Reveal (1993) = EricalesAdoxales Nakai (1949) = DipsacalesAesculales Bromhead (1838) = SapindalesAgavales Hutch (1934) = AsparagalesAkaniales Doweld (2001) = BrassicalesAlismatales Dumort (1829)Alliales Traub (1972) = Asparagales

422 AGP II


Alseuosmiales Doweld (2001) = AsteralesAlstroemeriales Hutch (1934) = LilialesAltingiales Doweld (1998) = SaxifragalesAmaranthales Dumort (1829) = CaryophyllalesAmaryllidales Bromhead (1840) = AsparagalesAmborellales Melikyan AVBobrov amp Zaytzeva

(1999) ndash family unplacedAmbrosiales Dumort (1829) = AsteralesAmmiales Small (1903) = ApialesAmomales Lindl (1835) = ZingiberalesAncistrocladales Takht ex Reveal (1992) =

CaryophyllalesAnisophylleales (Benth amp Hookf) Takht ex

Reveal amp Doweld (1999)Annonales Lindl (1833) = MagnolialesAnthobolales Dumort (1829) = SantalalesApiales Nakai (1930)Apocynales Bromhead (1838) = GentianalesAponogetonales Hutch (1934) = AlismatalesAquifoliales Senft (1856)Arales Dumort (1829) = AlismatalesAraliales Reveal (1996) = ApialesAralidiales Takht ex Reveal (1992) = ApialesArecales Bromhead (1840)Aristolochiales Dumort (1829) = PiperalesAsarales Horan (1847) = PiperalesAsclepiadales Dumort (1829) = GentianalesAsparagales Bromhead (1838)Asphodelales Doweld (2001) = AsparagalesAsteliales Dumort (1829) = AsparagalesAsterales Lindl (1833)Atriplicales Horan (1847) = CaryophyllalesAucubales Takht (1997) = GarryalesAustrobaileyales Takht ex Reveal (1992)Avenales Bromhead (1838) = PoalesBalanitales CYWu (2002) ndash family unplaced in

eurosids I = ZygophyllalesBalanopales Engl (1897) = MalpighialesBalanophorales Dumort (1829) ndash family unplaced

at end of systemBalsaminales Lindl (1833) = EricalesBarbeyales Takht amp Reveal (1993) = RosalesBarclayales Doweld (2001) = Nymphaeales family

unplaced at beginning of systemBatales Engl (1907) = BrassicalesBegoniales Dumort (1829) = CucurbitalesBerberidales Dumort (1829) = RanunculalesBerberidopsidales Doweld (2001) ndash family

unplaced in core eudicotsBetulales Bromhead (1838) = FagalesBiebersteiniales Takht (1997) = SapindalesBignoniales Lindl (1833) = LamialesBixales Lindl (1833) = MalvalesBoraginales Dumort (1829) ndash family unplaced

under euasterid IBrassicales Bromhead (1838)

Brexiales Lindl (1833) = CelastralesBromeliales Dumort (1829) = PoalesBruniales Dumort (1829) ndash family unplaced

under euasterid IIBrunoniales Lindl (1833) = AsteralesBurmanniales Heinze (1927) = DioscorealesBurserales Baskerville (1839) = SapindalesButomales Hutch (1934) = AlismatalesBuxales Takht ex Reveal (1996) ndash family

unplaced under eudicotsByblidales Nakai ex Reveal (1993) = LamialesCactales Dumort (1829) = CaryophyllalesCallitrichales Dumort (1829) = LamialesCalycanthales Mart (1835) = LauralesCalycerales Takht ex Reveal (1996) = AsteralesCampanulales Rchb (1828) = AsteralesCampynematales Doweld (2001) = LilialesCanellales Cronquist (1957)Cannales Dumort (1829) = ZingiberalesCapparales Hutch (1924) = BrassicalesCaprifoliales Lindl (1833) = DipsacalesCardiopteridales Takht (1997) = AquifolialesCarduales Small (1903) = AsteralesCaricales LDBenson (1957) = BrassicalesCarlemanniales Doweld (2001) = LamialesCaryophyllales Perleb (1826)Cassiales Horan (1847) = FabalesCasuarinales Lindl (1833) = FagalesCelastrales Baskerville (1839)Centrolepidales RDahlgren ex Takht (1997) =

PoalesCephalotales Nakai (1943) = OxalidalesCeratophyllales Bisch (1839)Cercidiphyllales Hu ex Reveal (1993) =

SaxifragalesChenopodiales Dumort (1829) = CaryophyllalesChironiales Griseb (1854) = GentianalesChloranthales ACSm ex J-FLeroy (1983) ndash

family unplaced at beginning of systemChrysobalanales (DC) Takht ex Reveal amp Dow-

eld (1999) = MalpighialesCinchonales Lindl (1835) = GentianalesCircaeasterales Takht (1997) = RanunculalesCistales Rchb (1828) = MalvalesCitrales Dumort (1829) = SapindalesCocosales Nakai (1930) = ArecalesColchicales Dumort (1829) = LilialesColumelliales Doweld (2001) ndash family unplaced in

euasterids IICombretales Baskerville (1839) = MyrtalesCommelinales Dumort (1829)Connarales Takht ex Reveal (1996) = OxalidalesConvolvulales Dumort (1829) = SolanalesCoriariales Lindl (1833) = CucurbitalesCornales Dumort (1829)Corylales Dumort (1829) = Fagales



Corynocarpales Takht (1997) = CucurbitalesCrassulales Lindl (1833) = SaxifragalesCrossosomatales Takht ex Reveal (1993)Cucurbitales Dumort (1829)Cunoniales Hutch (1924) = OxalidalesCyclanthales JHSchaffn (1911) = PandanalesCymodoceales Nakai (1943) = AlismatalesCynarales Raf (1837) = AsteralesCynomoriales Burnett (1835) ndash type genus

unplaced at end of systemCyperales Wettst (1911) = PoalesCyrillales Doweld (2001) = EricalesCytinales Dumort (1829) ndash type genus unplaced

at end of systemDaphnales Lindl (1833) = MalvalesDaphniphyllales Pulle ex Cronquist (1981) =

SaxifragalesDasypogonales Doweld (2001) ndash family unplaced

under commelinidsDatiscales Dumort (1829) = CucurbitalesDegeneriales CYWu (2002) = MagnolialesDesfontainiales Takht (1997) ndash family unplaced

under euasterids IIDiapensiales Engl amp Gilg (1924) = EricalesDidymelales Takht (1967) ndash see BuxalesDilleniales Hutch (1924) ndash family unplaced under

core eudicotsDioncophyllales Takht ex Reveal (1993) =

CaryophyllalesDioscoreales Hookf (1873)Diospyrales Prantl (1874) = EricalesDipentodontales CYWu (2002) ndash type genus

unplaced at end of systemDipsacales Dumort (1829)Droserales Griseb (1854) = CaryophyllalesEbenales Engl (1892) = EricalesEchiales Lindl (1838) ndash see BoraginalesElaeagnales Bromhead (1838) = RosalesElaeocarpales Takht (1997) = OxalidalesElatinales Nakai (1949) = MalpighialesElodeales Nakai (1950) = AlismatalesEmmotales Doweld (2001) = Icacinales unplaced

family under euasterids IEmpetrales Raf (1838) = EricalesEricales Dumort (1829)Eriocaulales Nakai (1930) = PoalesErythropalales Tiegh (1899) = SantalalesEscalloniales Doweld (2001) ndash family unplaced in

euasterids IIEucommiales Nemejc ex Cronquist (1981) =

GarryalesEuphorbiales Lindl (1833) = MalpighialesEupomatiales Takht ex Reveal (1992) =

MagnolialesEupteleales Hu ex Reveal (1993) =

Ranunculales

Euryalales HLLi (1955) ndash see NymphaealesFabales Bromhead (1838)Fagales Engl (1892)Ficales Dumort (1829) = RosalesFlacourtiales Heinze (1927) = MalpighialesFlagellariales (Meisn) Takht ex Reveal amp Dow-

eld (1999) = PoalesFouquieriales Takht ex Reveal (1992) = EricalesFrancoales Takht (1997) = GeranialesFrangulales Wirtg (1860) = RosalesGaliales Bromhead (1838) = GentianalesGarryales Lindl (1846)Geissolomatales Takht ex Reveal (1992) ndash family

unplaced under core eudicotsGentianales Lindl (1833)Geraniales Dumort (1829)Gesneriales Dumort (1829) = LamialesGlaucidiales Takht ex Reveal (1992) =

RanunculalesGlobulariales Dumort (1829) = LamialesGoodeniales Lindl (1833) = AsteralesGreyiales Takht (1997) = GeranialesGriseliniales (JRForst amp GForst ex ACunn)

Takht ex Reveal amp Doweld (1999) = ApialesGrossulariales Lindl (1833) = SaxifragalesGrubbiales Doweld (2001) = CornalesGunnerales Takht ex Reveal (1992)Gyrocarpales Dumort (1829) = LauralesGyrostemonales Takht (1997) = BrassicalesHaemodorales Hutch (1934) = CommelinalesHaloragales Bromhead (1838) = SaxifragalesHamamelidales Griseb (1854) = SaxifragalesHanguanales RDahlgren ex Reveal (1992) =

CommelinalesHeisteriales Tiegh (1899) = SantalalesHelleborales Nakai (1949) = RanunculalesHelwingiales Takht (1997) = AquifolialesHimantandrales Doweld amp Shevyryova (1998) =

MagnolialesHippuridales Thomeacute (1874) = LamialesHomaliales Bromhead (1838) = MalpighialesHortensiales Griseb (1854) = CornalesHuales Doweld (2001) = MalpighialesHuerteales Doweld (2001) ndash see Tapisciaceae an

unplaced family in rosidsHydatellales (UHamann) Cronquist ex Reveal amp

Doweld (1999) = PoalesHydnorales Takht ex Reveal (1992) = PiperalesHydrangeales Nakai (1943) = CornalesHydrastidales Takht (1997) = RanunculalesHydrocharitales Dumort (1829) = AlismatalesHydropeltidales Spenn (1834) ndash see

NymphaeaceaeHydrostachyales Diels ex Reveal (1993) =

CornalesHypericales Dumort (1829) = Malpighiales

424 AGP II


Hypoxidales Takht ex Reveal amp Doweld (1999) =Asparagales

Icacinales Tiegh (1899) ndash family unplaced undereuasterids I

Illiciales Hu ex Cronquist (1981) = Austrobailey-ales

Iridales Raf (1815) = AsparagalesIrvingiales Doweld (2001) = MalpighialesIteales Doweld (2001) = SaxifragalesIxerbales Doweld (2001) ndash family unplaced in

rosidsIxiales Lindl (1835) = AsparagalesJasminales Dumort (1829) = LamialesJuglandales Dumort (1829) = FagalesJulianiales Engl (1907) = SapindalesJuncaginales Hutch (1934) = AlismatalesJuncales Dumort (1829) = PoalesLacistematales Baskerville (1839) = MalpighialesLactoridales Takht ex Reveal (1993) =

PiperalesLamiales Bromhead (1838)Lardizabalales Loconte (1995) = RanunculalesLaurales Perleb (1826)Lecythidales Cronquist (1957) = EricalesLedocarpales Doweld (2001) = GeranialesLeitneriales Engl (1897) = SapindalesLentibulariales Lindl (1833) = LamialesLigustrales Bartl ex Bisch (1839) = LamialesLiliales Perleb (1826)Limnanthales Nakai (1930) = BrassicalesLinales Baskerville (1839) = MalpighialesLoasales Bessey (1907) = CornalesLobeliales Drude (1888) = AsteralesLoganiales Lindl (1833) = GentianalesLonicerales TLiebe (1866) = DipsacalesLoranthales Dumort (1829) = SantalalesLowiales Takht ex Reveal amp Doweld (1999) =

ZingiberalesLythrales Caruel (1881) = MyrtalesMagnoliales Bromhead (1838)Malpighiales Mart (1835)Malvales Dumort (1829)Marathrales Dumort (1829) = MalpighialesMarcgraviales Doweld (2001) = EricalesMayacales Nakai (1943) = PoalesMedusagynales Takht ex Reveal amp Doweld

(1999) = MalpighialesMedusandrales Brenan (1952) ndash type genus

unplaced at end of systemMelanthiales RDahlgren ex Reveal (1992) =

LilialesMelastomatales Oliv (1895) = MyrtalesMeliales Lindl (1833) = SapindalesMelianthales Doweld = GeranialesMeliosmales CYWu (2002) ndash see SabialesMenispermales Bromhead (1838) = Ranunculales

Menyanthales TYamaz ex Takht (1997) =Asterales

Metteniusales Takht (1997) ndash type genusunplaced at end of system

Miyoshiales Nakai (1941) ndash see Petrosavialesfamily unplaced under monocots

Monimiales Dumort (1829) = LauralesMoringales Nakai (1943) = BrassicalesMusales Reveal (1997) = ZingiberalesMyricales Engl (1897) = FagalesMyristicales Thomeacute (1877) = MagnolialesMyrothamnales Nakai ex Reveal (1993) =

GunneralesMyrsinales Spenn (1835) = EricalesMyrtales Rchb (1828)Najadales Dumort (1829) = AlismatalesNandinales Doweld (2001) = RanunculalesNarcissales Dumort (1829) = AsparagalesNartheciales Reveal amp Zomlefer (1998) =

DioscorealesNelumbonales Willk amp Lange (1861) = ProtealesNepenthales Dumort (1829) = CaryophyllalesNeuradales Doweld (2001) = MalvalesNitrariales Doweld (2001) = SapindalesNolanales Lindl (1835) = SolanalesNothofagales Doweld (2001) = FagalesNyctaginales Dumort (1829) = CaryophyllalesNymphaeales Dumort (1829) = family unplaced

at beginning of systemOchnales Hutch ex Reveal (1992) = MalpighialesOenotherales Bromhead (1838) = MyrtalesOlacales Benth amp Hookf (1862) = SantalalesOleales Lindl (1833) = LamialesOnagrales Rchb (1828) = MyrtalesOncothecales Doweld (2001) ndash family unplaced

under euasterids IOpuntiales Endl ex Willk (1854) =

CaryophyllalesOrchidales Raf (1815) = AsparagalesOxalidales Heintze (1927)Paeoniales Heinze (1927) = SaxifragalesPandales Engl amp Gilg (1912ndash13) = MalpighialesPandanales Lindl (1833)Papaverales Dumort (1829) = RanunculalesParacryphiales Takht ex Reveal (1992) ndash family

unplaced under euasterid IIParidales Dumort (1829) = LilialesParnassiales Nakai (1943) = CelastralesPassiflorales Dumort (1829) = MalpighialesPenaeales Lindl (1833) = MyrtalesPennantiales Doweld (2001) = ApialesPentaphragmatales Doweld (2001) = AsteralesPetiveriales Lindl (1833) = CaryophyllalesPetrosaviales Takht (1997) ndash family unplaced

under monocotsPhellinales Doweld (2001) = Asterales



Philydrales Dumort (1829) = CommelinalesPhyllanthales Doweld (2001) = MalpighialesPhysenales Takht (1977) = CaryophyllalesPhytolaccales Doweld (2001) = CaryophyllalesPicramniales Doweld (2001) ndash family unplaced

under rosidsPinguiculales Dumort (1829) = LamialesPiperales Dumort (1829)Pittosporales Lindl (1833) = ApialesPlantaginales Lindl (1833) = LamialesPlatanales JHSchaffn (1911) = ProtealesPlumbaginales Lindl (1833) = CaryophyllalesPoales Small (1903)Podophyllales Dumort (1829) = RanunculalesPodostemales Lindl (1833) = MalpighialesPolemoniales Bromhead (1838) = EricalesPolygalales Dumort (1829) = FabalesPolygonales Dumort (1829) = CaryophyllalesPontederiales Hookf (1873) = CommelinalesPortulacales Dumort (1829) = CaryophyllalesPosidoniales Nakai (1943) = AlismatalesPotamogetonales Dumort (1829) = AlismatalesPrimulales Dumort (1829) = EricalesProteales Dumort (1829)Quercales Burnett (1835) = FagalesQuillajales Doweld (2001) = FabalesQuintiniales Doweld (2001) = Sphenostemonales

unplaced under euasterids IIRafflesiales Oliv (1895) ndash unplaced family type at

end of systemRanunculales Dumort (1829)Rapateales (Meisn) Colella ex Reveal amp Doweld

= PoalesResedales Dumort (1829) = BrassicalesRestionales Hookf (1873) = PoalesRhabdodendrales Doweld (2001) = CaryophyllalesRhamnales Dumort (1829) = RosalesRhinanthales Dumort (1829) = LamialesRhizophorales (Pers) Reveal amp Doweld (1999) =

MalpighialesRhodorales Horan (1847) = EricalesRhoipteleales Novaacutek ex Reveal (1992) = FagalesRoridulales Nakai (1943) = EricalesRosales Perleb (1826)Rousseales Doweld (2001) = AsteralesRubiales Dumort (1829) = GentianalesRuppiales Nakai (1950) = AlismatalesRutales Perleb (1826) = SapindalesSabiales Takht (1987) = family unplaced under

eudicotsSalicales Lindl (1833) = MalpighialesSalvadorales RDahlgren ex Reveal (1993) =

BrassicalesSamolales Dumort (1829) = EricalesSamydales Dumort (1829) = MalpighialesSanguisorbales Dumort (1829) = Rosales

Santalales Dumort (1829)Sapindales Dumort (1829)Sapotales Hookf (1868) = EricalesSarraceniales Bromhead (1838) = EricalesSaxifragales Dumort (1829)Scheuchzeriales BBoivin (1956) = AlismatalesScleranthales Dumort (1829) = CaryophyllalesScrophulariales Lindl (1833) = LamialesScyphostegiales Croizat (1994) = MalpighialesSedales Rchb (1828) = SaxifragalesSilenales Lindl (1833) = CaryophyllalesSimmondsiales Reveal (1992) = CaryophyllalesSmilacales Lindl (1833) = LilialesSolanales Dumort (1829)Sphenocleales Doweld (2001) = SolanalesSphenostemonales Doweld (2001) ndash family

unplaced under euasterids IIStellariales Dumort (1829) = CaryophyllalesStemonales Takht ex Doweld (2001) =

PandanalesStilbales Doweld (2001) = LamialesStylidiales Takht ex Reveal (1992) = AsteralesStyracales Bisch (1839) = EricalesSurianales Doweld (2001) = FabalesTaccales Dumort (1829) = DioscorealesTamales Dumort (1829) = DioscorealesTamaricales Hutch (1924) = CaryophyllalesTecophilaeales Traub ex Reveal (1993) =

AsparagalesTernstroemiales Doweld (2001) = EricalesTheales Lindl (1833) = EricalesTheligonales Nakai (1942) = GentianalesThymelaeales Willk (1854) = MalvalesTiliales Caruel (1881) = MalvalesTofieldiales Reveal amp Zomlefer (1998) =

AlismatalesTorricelliales Takht ex Reveal amp Doweld (1999) =

ApialesTovariales Nakai (1943) = BrassicalesTribelales Doweld (2001) ndash family unplaced in

euasterids IITrilliales Takht (1997) = LilialesTrimeniales Doweld (2001) = AustrobaileyalesTriuridales Hookf (1873) = PandanalesTrochodendrales Takht ex Cronquist (1981) ndash

unplaced family under eudicotsTropaeolales Takht ex Reveal (1992) =

BrassicalesTurnerales Dumort (1829) = MalpighialesTyphales Dumort (1829) = PoalesUlmales Lindl (1833) = RosalesUrticales Dumort (1829) = RosalesVacciniales Dumort (1829) = EricalesVahliales Doweld (2001) ndash family unplaced in

euasterids IVallisneriales Nakai (1949) = Alismatales

426 AGP II


Velloziales RDahlgren ex Reveal (1992) =Pandanales

Veratrales Dumort (1829) = LilialesVerbenales Horan (1847) = LamialesViburnales Dumort (1829) = DipsacalesVincales Horan (1847) = GentianalesViolales Perleb (1826) = MalpighialesViscales Tiegh (1899) = SantalalesVitales Reveal (1996) ndash family unplaced under

core eudicotsVochysiales Dumort (1829) = MyrtalesWinterales (Meisn) AC Sm ex Reveal (1993) =

CanellalesXanthorrhoeales Takht ex Reveal amp Doweld

(1999) = AsparagalesXimeniales Tiegh (1899) = SantalalesXyridales Lindl (1846) = PoalesZingiberales Griseb (1854)Zosterales Nakai (1943) = AlismatalesZygophyllales Chalk (1990) ndash family unplaced

under eurosid I

SELECTED FAMILIAL SYNONYMS

The following names are primarily those in currentuse or listed here so as to define more clearly therecognized families Accepted family names are inbold face Families included as belonging to typegenera of an uncertain position are in italics

Abolbodaceae Nakai (1943) = XyridaceaeAbrophyllaceae Nakai (1943) = RousseaceaeAcanthaceae Juss (1789) nom consAcanthochlamydaceae PCKao (1989) =

VelloziaceaeAceraceae Juss (1789) nom cons = SapindaceaeAchariaceae Harms (1897) nom consAchatocarpaceae Heimerl (1934) nom consAchradaceae Vest (1818) = SapotaceaeAcoraceae Martynov (1820)Actinidiaceae Gilg amp Werderm (1825) nom

consAdoxaceae EMey (1839) nom consAegialitidaceae Lincz (1968) = PlumbaginaceaeAegicerataceae Blume (1833) = MyrsinaceaeAextoxicaceae Engl amp Gilg (1920) nom consAgapanthaceae FVoigt (1850) optional syn-

onym of AlliaceaeAgavaceae Dumort (1829) nom cons optional

synonym of AsparagaceaeAgdestidaceae Nakai (1942) = PhytolaccaceaeAizoaceae Martynov (1820) nom consAkaniaceae Stapf (1912) nom consAlangiaceae DC (1827) nom cons = CornaceaeAldrovandaceae Nakai (1949) = DroseraceaeAlismataceae Vent (1799) nom cons

Alliaceae Batsch ex Borkh (1797) nom consAloaceae Batsch (1802) = Asphodelaceae optional

synonym of XanthorrhoeaceaeAlseuosmiaceae Airy Shaw (1964)Alsinaceae Bartl (1825) nom cons =

CaryophyllaceaeAlstroemeriaceae Dumort (1829) nom consAltingiaceae Horan (1843) nom consAlzateaceae SAGraham (1985)Amaranthaceae Juss (1789) nom consAmaryllidaceae JSt-Hil (1805) nom cons

optional synonym of AlliaceaeAmborellaceae Pichon (1948) nom consAmbrosiaceae Martynov (1820) nom cons =

AsteraceaeAmygdalaceae Marquis (1820) nom cons =

RosaceaeAmyridaceae Kunth (1824) = RutaceaeAnacardiaceae RBr (1818) nom consAnarthriaceae DFCutler amp Airy Shaw (1965)Ancistrocladaceae Planch ex Walp (1851)

nom consAndrostachyaceae Airy Shaw (1964) =

PicrodendraceaeAnemarrhenaceae Conran MWChase amp Rudall

(1997) = Agavaceae optional synonym ofAsparagaceae

Anisophylleaceae Ridl (1922)Annonaceae Juss (1789) nom consAnomochloaceae Nakai (1943) = PoaceaeAnopteraceae Doweld (2001) = EscalloniaceaeAnthericaceae JAgardh (1858) = Agavaceae

optional synonym of AsparagaceaeAntirrhinaceae Pers (1807) = PlantaginaceaeAntoniaceae Hutch (1959) = LoganiaceaeAphanopetalaceae Doweld (2001)Aphloiaceae Takht (1985)Aphyllanthaceae Burnett (1835) optional syn-

onym of AsparagaceaeApiaceae Lindl (1836) nom consApocynaceae Juss (1789) nom consApodanthaceae (RBr) Tiegh ex Takht (1987) =

RafflesiaceaeAponogetonaceae JAgardh (1858) nom consApostasiaceae Lindl (1833) nom cons =

OrchidaceaeAptandraceae Miers (1853) = OlacaceaeAquifoliaceae DC ex ARich (1828) nom consAquilariaceae RBr ex DC (1825) =

ThymelaeaceaeAraceae Juss (1789) nom consAragoaceae DDon (1835) = PlantaginaceaeAraliaceae Juss (1789) nom consAralidiaceae Philipson amp BCStone (1980)Arecaceae Schultz-Sch (1832) nom consArgophyllaceae (Engl) Takht 1987



Aristoteliaceae Dumort (1829) = ElaeocarpaceaeAristolochiaceae Juss (1789) nom consAsclepiadaceae Borkh (1797) nom cons =

ApocynaceaeAsparagaceae Juss (1789) nom consAsphodelaceae Juss (1789) optional synonym

of XanthorrhoeaceaeAspidistraceae Endl (1841) = Ruscaceae optional

synonym of AsparagaceaeAsteliaceae Dumort (1829)Asteraceae Martynov (1820) nom consAsteranthaceae RKnuth (1939) nom cons =

LecythidaceaeAsteropeiaceae (Szyszyl) Takht ex Reveal amp

Hoogland (1990)Atherospermataceae RBr (1814)Aucubaceae JAgardh (1858) optional synonym

of GarryaceaeAustrobaileyaceae (Croizat) Croizat 1943 nom

consAverrhoaceae Hutch (1959) = OxalidaceaeAvetraceae Takht (1997) = DioscoreaceaeAvicenniaceae Endl (1841) = AcanthaceaeBalanitaceae Endl (1841) nom cons =

ZygophyllaceaeBalanitaceae Endl (1841) = ZygophyllaceaeBalanopaceae Benth amp Hookf (1880) nom

consBalanophoraceae Rich (1822) nom cons

unplacedBalsaminaceae Bercht amp JPresl (1820) nom

consBambusaceae Burnett (1835) = PoaceaeBarbeuiaceae Nakai (1942)Barbeyaceae Rendle (1916) nom consBarclayaceae HLLi (1955) = NymphaeaceaeBarringtoniaceae FRudolphi (1830) nom cons =

LecythidaceaeBasellaceae Raf (1837) nom consBataceae Perleb (1838) nom consBaueraceae Lindl (1830) = CunoniaceaeBaxteriaceae Takht (1996) = DasypogonaceaeBegoniaceae Bercht amp JPresl (1820) nom

consBehniaceae Conran MWChase amp Rudall (1997)

= Agavaceae optional synonym of AsparagaceaeBembiciaceae RCKeating amp Takht (1996) =

SalicaceaeBerberidaceae Juss (1789) nom consBerberidopsidaceae Takht (1985)Berryaceae Doweld (2001) = MalvaceaeBersamaceae Doweld = MelianthaceaeBerzeliaceae Nakai (1943) = BruniaceaeBetulaceae Gray (1821) nom consBiebersteiniaceae Endl (1841)Bignoniaceae Juss (1789) nom cons

Bischofiaceae Airy Shaw (1964) = PhyllanthaceaeBixaceae Kunth (1822) nom consBlandfordiaceae RDahlgren amp Clifford

(1985)Blepharocaryaceae Airy Shaw (1964) =

AnacardiaceaeBoerlagellaceae HJLam (1925) = SapotaceaeBombacaceae Kunth (1822) nom cons = Mal-

vaceaeBonnetiaceae (Bartl) LBeauv ex Nakai (1948)Boopidaceae Cass (1816) = CalyceraceaeBoraginaceae Juss (1789) nom consBoryaceae (Baker) MWChase Rudall amp Conran

(1997)Brassicaceae Burnett (1835) nom consBretschneideraceae Engl amp Gilg (1924) nom

cons optional synonym of AkaniaceaeBrexiaceae Loudon (1830) = CelastraceaeBromeliaceae Juss (1789) nom consBrunelliaceae Engl (1897) nom consBruniaceae Bercht amp JPresl (1820) nom consBrunoniaceae Dumort (1829) nom cons =

GoodeniaceaeBuddlejaceae KWilh (1910) nom cons =

ScrophulariaceaeBurchardiaceae Takht (1996) = ColchicaceaeBurmanniaceae Blume (1827) nom consBurseraceae Kunth (1824) nom consButomaceae Mirb (1804) nom consBuxaceae Dumort (1822) nom consByblidaceae (Engl amp Gilg) Domin 1922 nom

consByttneriaceae RBr (1814) nom cons =

MalvaceaeCabombaceae Rich ex ARich (1822) nom

cons optional synonym of NymphaeaceaeCactaceae Juss (1789) nom consCaesalpiniaceae RBr (1814) nom cons =

FabaceaeCalceolariaceae (DDon) Olmstead (2001)Calectasiaceae Endl (1838) = DasypogonaceaeCalligonaceae Chalk (1985) = PolygonaceaeCallitrichaceae Bercht amp JPresl (1820) nom

cons = PlantaginaceaeCalochortaceae Dumort (1829) = LiliaceaeCalycanthaceae Lindl (1819) nom consCalyceraceae RBr ex Rich (1820) nom

consCampanulaceae Juss (1789) nom consCampynemataceae Dumort (1829)Canacomyricaceae Baum-Bod ex Doweld (2001)

= MyricaceaeCanellaceae Mart (1832) nom consCannabaceae Martynov (1820) nom consCannaceae Juss (1789) nom consCanotiaceae Airy Shaw (1964) = Celastraceae

428 AGP II


Capparaceae Juss (1789) nom cons =Brassicaceae

Caprifoliaceae Juss (1789) nom consCardiopteridaceae Blume (1847) nom consCaricaceae Dumort (1829) nom consCarlemanniaceae Airy Shaw (1964)Carpinaceae Vest (1818) = BetulaceaeCarpodetaceae Fenzl (1841) = RousseaceaeCartonemataceae Pichon (1946) = CommelinaceaeCaryocaraceae Voigt (1845) nom consCaryophyllaceae Juss (1789) nom consCassythaceae Bartl ex Lindl (1833) nom cons =

LauraceaeCasuarinaceae RBr (1814) nom consCecropiaceae CCBerg (1978) = UrticacaeaeCelastraceae RBr (1814) nom consCeltidaceae Link (1831) nom cons = Canna-

baceaeCentrolepidaceae Endl (1836) nom consCephalotaceae Dumort (1829) nom consCeratophyllaceae Gray (1821) nom consCercidiphyllaceae Engl (1907) nom consChenopodiaceae Vent (1799) nom cons =

AmaranthaceaeChionographidaceae Takht (1966) =

MelanthiaceaeChloanthaceae Hutch (1959) = LamiaceaeChloranthaceae RBr ex Sims (1820) nom

consChrysobalanaceae RBr (1818) nom consCichoriaceae Juss (1789) nom cons = AsteraceaeCircaeasteraceae Hutch (1926) nom consCistaceae Juss (1789) nom consCleomaceae Horan (1834) = BrassicaceaeClethraceae Klotzsch (1851) nom consClusiaceae Lindl (1836) nom consCneoraceae Vest (1818) nom cons = RutaceaeCobaeaceae DDon (1824) = PolemoniaceaeCochlospermaceae Planch (1847) nom cons

optional synonym of BixaceaeColchicaceae DC (1804) nom consColumelliaceae DDon (1828) nom consCombretaceae RBr (1810) nom consCommelinaceae Mirb (1804) nom consCompositae Giseke (1792) nom alt et cons =

AsteraceaeConnaraceae RBr (1818) nom consConostylidaceae (Benth) Takht (1987) =

HaemodoraceaeConvallariaceae Horan (1834) = Ruscaceae

optional synonym of AsparagaceaeConvolvulaceae Juss (1789) nom consCordiaceae RBr ex Dumort (1829) nom cons =

BoraginaceaeCoriariaceae DC (1824) nom consCoridaceae JAgardh (1858) = Myrsinaceae

Cornaceae Dumort (1829) nom consCorokiaceae Kapil ex Takht (1997) =

ArgophyllaceaeCorsiaceae Becc (1878) nom consCorylaceae Mirb (1815) nom cons = BetulaceaeCorynocarpaceae Engl (1897) nom consCostaceae Nakai (1941)Crassulaceae JSt-Hil (1805) nom consCroomiaceae Nakai (193) = StemonaceaeCrossosomataceae Engl (1897) nom consCruciferae Juss (1789) nom alt et cons =

BrassicaceaeCrypteroniaceae ADC (1868) nom consCtenolophonaceae (HWinkl) Exell amp

Mendonccedila (1951)Cucurbitaceae Juss (1789) nom consCunoniaceae RBr (1814) nom consCurtisiaceae (Engl) Takht (1987)Cuscutaceae Bercht amp JPresl (1820) nom cons

= ConvolvulaceaeCyananthaceae JAgardh (1858) =

CampanulaceaeCyanastraceae Engl (1900) nom cons =

TecophilaeaceaeCyclanthaceae Poit ex ARich (1824) nom

consCyclocheilaceae Marais (1981) = OrobanchaceaeCymodoceaceae NTaylor (1909) nom consCynomoriaceae Lindl (1833) nom cons

unplacedCyperaceae Juss (1789) nom consCyphiaceae ADC (1839) = Lobeliaceae optional

synonym of CampanulaceaeCyphocarpaceae (Miers) Reveal amp Hoogl (1996) =

Lobeliaceae optional synonym of Campanu-laceae

Cypripediaceae Lindl (1833) = OrchidaceaeCyrillaceae Endl (1841) nom consCytinaceae ARich (1824) unplacedDactylanthaceae (Engl) Takht (1987) =

BalanophoraceaeDaphniphyllaceae Muumlll-Arg (1869) nom consDasypogonaceae Dumort (1829)Datiscaceae Bercht amp JPresl (1820) nom consDavidiaceae HLLi (1955) = CornaceaeDavidsoniaceae Bange (1952) = CunoniaceaeDecaisneaceae (Takht ex H N Qin) Loconte

(1995) = LardizabalaceaeDegeneriaceae IWBailey amp ACSm (1942)

nom consDesfontainiaceae Endl (1841) nom cons

optional synonym of ColumelliacaeDialypetalanthaceae Rizzini amp Occhioni (1948)

nom cons = RubiaceaeDianellaceae Salisb (1866) = Hemerocallidaceae

optional synonym of Xanthorrhoeaceae



Diapensiaceae Lindl (1836) nom consDichapetalaceae Baill (1886) nom cons

optional synonym of ChrysobalanaceaeDichondraceae Dumort (1829) = ConvolvulaceaeDiclidantheraceae J Agardh (1858) nom cons =

PolygalaceaeDidiereaceae Radlk (1896) nom consDidymelaceae Leandri (1937) optional syn-

onym of BuxaceaeDiegodendraceae Capuron (1964) optional syn-

onym of BixaceaeDiervillaceae (Raf) Pyck (1998) optional syn-

onym of CaprifoliaceaeDilleniaceae Salisb (1807) nom consDionaeaceae Raf (1837) = DroseraceaeDioncophyllaceae Airy Shaw (1952) nom consDioscoreaceae RBr (1810) nom consDipentodontaceae Merr (1941) nom cons

unplacedDipsacaceae Juss (1789) nom cons optional

synonym of CaprifoliaceaeDipterocarpaceae Blume (1825) nom consDirachmaceae Hutch (1959)Donatiaceae BChandler (1911) nom cons

optional synonym of StylidiaceaeDoryanthaceae RDahlgren amp Clifford (1985)Dracaenaceae Salisb (1866) = Ruscaceae

optional synonym of AsparagaceaeDroseraceae Salisb (1808) nom consDrosophyllaceae Chrtek Slaviacutekovaacute amp Stud-

nicka (1989)Duabangaceae Takht (1986) = LythraceaeDuckeodendraceae Kuhlm (1950) = SolanaceaeDysphaniaceae (Pax) Pax (1927) nom cons =

AmaranthaceaeEbenaceae Guumlrke (1891) nom consEcdeiocoleaceae DFCutler amp Airy Shaw (1965)Ehretiaceae Mart (1827) nom cons =

BoraginaceaeElaeagnaceae Juss (1789) nom consElaeocarpaceae Juss ex DC (1816) nom consElatinaceae Dumort (1829) nom consEllisiophyllaceae Honda (1930) = PlantaginaceaeEmblingiaceae Airy Shaw (1964)Emottaceae Tiegh (1899) = IcacinaceaeEmpetraceae Bercht amp JPresl (1820) nom cons

= EricaceaeEngelhardtiaceae Reveal amp Doweld (1999) =

JuglandaceaeEpacridaceae RBr (1810) nom cons = EricaceaeEpimediaceae Menge (1839) = BerberidaceaeEremolepidaceae Tiegh ex Nakai (1952) =

SantalaceaeEremosynaceae Dandy (1959)Ericaceae Juss (1789) nom consEriocaulaceae Martynov (1820) nom cons

Eriospermaceae Endl (1841) = Ruscaceaeoptional synonym of Asparagaceae

Erycibaceae Endl ex Meisn (1840) =Convolvulaceae

Erythropalaceae Pilg amp KKrause (1914) nomcons = Olacaceae

Erythroxylaceae Kunth (1822) nom consEscalloniaceae RBr ex Dumort (1829) nom

consEschscholziaceae Ser (1847) = PapaveraceaeEucommiaceae Engl (1909) nom consEucryphiaceae Endl (1841) nom cons =

CunoniaceaeEuphorbiaceae Juss (1789) nom consEuphroniaceae Marc-Berti (1989) optional

synonym of ChrysobalanaceaeEupomatiaceae Endl (1841) nom consEupteleaceae KWilh (1910) nom consEuryalaceae JAgardh (1858) = NymphaeaceaeEustrephaceae Chupov (1994) = Laxmanniaceae

optional synonym of AsparagaceaeExbucklandiaceae Reveal amp Doweld (1999) =

HamamelidaceaeExocarpaceae JAgardh (1858) = SantalaceaeFabaceae Lindl (1836) nom consFagaceae Dumort (1829) nom consFlacourtiaceae Rich (1815-1816) nom cons =

SalicaceaeFlagellariaceae Dumort (1829) nom consFlindersiaceae CTWhite ex Airy Shaw (1964) =

RutaceaeFoetidiaceae Airy Shaw (1964) = LecythidaceaeFouquieriaceae DC (1828) nom consFrancoaceae AJuss (1832) nom cons optional

synonym of MelianthaceaeFrangulaceae DC (1805) = RhamnaceaeFrankeniaceae Desv (1817) nom consFumariaceae Bercht amp JPresl (1820) nom

cons optional synonym of PapaveraceaeGarryaceae Lindl (1834) nom consGeissolomataceae Endl (1841)Geitonoplesiaceae RDahlgren ex Conran (1994)

= Hemerocallidaceae optional synonym ofXanthorrhoeaceae

Gelsemiaceae (GDon) Struwe amp VAAlbert(1995)

Geniostomaceae Struwe amp VAAlbert (1995) =Loganiaceae

Gentianaceae Juss (1789) nom consGeosiridaceae Jonker (1939) nom cons =

IridaceaeGeraniaceae Juss (1789) nom consGesneriaceae Rich amp Juss ex DC (1816) nom

consGisekiaceae Nakai (1942)Glaucidiaceae Tamura (1972) = Ranunculaceae

430 AGP II


Globulariaceae DC (1805) nom cons =Plantaginaceae

Goetzeaceae Miers ex Airy Shaw (1964) =Solanaceae

Gomortegaceae Reiche (1896) nom consGonystylaceae Tiegh (1896) nom cons =

ThymelaeaceaeGoodeniaceae RBr (1810) nom consGoupiaceae Miers (1862)Gramineae Juss (1789) nom alt et cons =

PoaceaeGreyiaceae Hutch (1926) nom cons =

MelianthaceaeGriseliniaceae JRForst amp GForst ex ACunn

(1839)Gronoviaceae Endl (1841) = LoasaceaeGrossulariaceae DC (1805) nom consGrubbiaceae Endl (1839) nom consGunneraceae Meisn (1842) nom consGustaviaceae Burnett (1835) = LecythidaceaeGuttiferae Juss (1789) nom alt et cons =

ClusiaceaeGyrocarpaceae Dumort (1829) = HernandiaceaeGyrostemonaceae Endl (1841) nom consHachetteaceae Doweld (2001) = Balanophora-

ceaeHaemodoraceae RBr (1810) nom consHalesiaceae DDon (1828) = StyracaceaeHalophilaceae JAgardh (1858) = Hydrocharita-

ceaeHalophytaceae ASoriano (1984)Haloragaceae RBr (1814) nom consHamamelidaceae RBr (1818) nom consHanguanaceae Airy Shaw (1964)Hectorellaceae Philipson amp Skipw (1961) =

PortulacaceaeHeliamphoraceae Chrtek Slaviacutekovaacute amp Studnicka

(1992) = SarraceniaceaeHeliconiaceae Nakai (1941)Heliotropiaceae Schrad (1819) nom cons =

BoraginaceaeHelleboraceae Vest (1818) = RanunculaceaeHeloniadaceae JAgardh (1858) = MelanthiaceaeHelosaceae (Schott amp Endl) Bromhead (1840) =

BalanophoraceaeHelwingiaceae Decne (1836)Hemerocallidaceae RBr (1810) optional syn-

onym of XanthorrhoeaceaeHemimeridaceae Doweld (2001) = PlantaginaceaeHenriqueziaceae Bremek (1957) = RubiaceaeHernandiaceae Bercht amp JPresl (1820) nom

consHerreriaceae Endl (1841) = Agavaceae optional

synonym of AsparagaceaeHesperocallidaceae Traub (1972) optional syn-

onym of Asparagaceae

Heteropyxidaceae Engl amp Gilg (1920) nomcons

Himantandraceae Diels (1917) nom consHippocastanaceae ARich (1823) nom cons =

SapindaceaeHippocrateaceae Juss (1811) nom cons =

CelastraceaeHippuridaceae Vest (1818) nom cons =

PlantaginaceaeHopkinsiaceae BGBriggs amp LASJohnson

(2000) = AnarthriaceaeHoplestigmataceae Gilg (1924) nom cons

unplacedHortoniaceae (JRPerkins amp Gilg) ACSm (1971)

= MonimiaceaeHostaceae BMathew (1988) = Agavaceae

optional synonym of AsparagaceaeHuaceae AChev (1947)Huerteaceae Doweld (2001) = TapisciaceaeHugoniaceae Arn (1834) = LinaceaeHumbertiaceae Pichon (1947) nom cons =

ConvolvulaceaeHumiriaceae AJuss (1829) nom consHyacinthaceae Batsch ex Borkh (1797)

optional synonym of AsparagaceaeHydatellaceae UHamann (1976)Hydnoraceae CAgardh (1821) nom consHydrangeaceae Dumort (1829) nom consHydrastidaceae Martynov (1820) =

RanunculaceaeHydrocharitaceae Juss (1789) nom consHydrocotylaceae (Link) NHyl (1945) nom cons

= AraliaceaeHydroleaceae Bercht amp JPresl (1820)Hydropeltidaceae (DC) Dumort (1822) =

NymphaeaceaeHydrophyllaceae RBr (1817) nom cons =

BoraginaceaeHydrostachyaceae (Tul) Engl (1894) nom

consHymenocardiaceae Airy Shaw (1964) =

PhyllanthaceaeHypecoaceae Willk amp Lange (1880) =

PapaveraceaeHypericaceae Juss (1789) nom consHypoxidaceae RBr (1814) nom consHypseocharitaceae Wedd (1861) optional syn-

onym of GeraniaceaeIcacinaceae (Benth) Miers (1851) nom consIdiospermaceae STBlake (1972) =

CalycanthaceaeIllecebraceae RBr (1810) nom cons =

CaryophyllaceaeIlliciaceae ACSm (1947) nom cons optional

synonym of SchisandraceaeIridaceae Juss (1789) nom cons



Irvingiaceae (Engl) Exell amp Mendonccedila (1951)nom cons

Isophysidaceae (Hutch) FABarkley (1948) =Iridaceae

Iteaceae JAgardh (1858) nom consIxerbaceae Griseb (1854)Ixioliriaceae Nakai (1943)Ixonanthaceae Planch ex Miq (1858) nom

consJaponoliriaceae Takht (1996) = PetrosaviaceaeJohnsoniaceae Lotsy (1911) = Hemerocallidaceae

optional synonym of XanthorrhoeaceaeJoinvilleaceae Toml amp ACSm (1970)Juglandaceae DC ex Perleb (1818) nom consJulianiaceae Hemsl (1906) nom cons =

AnacardiaceaeJuncaceae Juss (1789) nom consJuncaginaceae Rich (1808) nom consJusticiaceae Raf (1838) = AcanthaceaeKaliphoraceae Takht (1996) = MontiniaceaeKiggelariaceae Link (1831) = AchariaceaeKingdoniaceae ASFoster ex Airy Shaw (1964)

optional synonym of CircaeasteraceaeKirengeshomaceae Nakai (1943) =

HydrangeaceaeKirkiaceae (Engl) Takht (1967)Koeberliniaceae Engl (1895) nom consKrameriaceae Dumort (1829) nom cons

optional synonym of ZygophyllaceaeLabiatae Juss (1789) nom alt et cons =

LamiaceaeLacandoniaceae EMartiacutenes amp Ramos (1989) =

TriuridaceaeLacistemataceae Mart (1826) nom consLactoridaceae Engl (1888) nom consLamiaceae Martynov (1820) nom consLanariaceae HHuber ex RDahlgren amp

AEvanWyk (1988)Langsdorffiaceae Tiegh ex Pilger (1914) =

BalanophoraceaeLardizabalaceae RBr (1821) nom consLauraceae Juss (1789) nom consLaxmanniaceae Bubani (1901-1902) optional

synonym of AsparagaceaeLecythidaceae ARich (1825) nom consLedocarpaceae Meyen (1834)Leeaceae Dumort (1829) nom cons = VitaceaeLeguminosae Juss (1789) nom alt et cons =

FabaceaeLeitneriaceae Benth amp Hookf (1880) nom cons

= SimaroubaceaeLemnaceae Martynov (1820) nom cons =

AraceaeLennoaceae Solms (1870) nom cons =

BoraginaceaeLentibulariaceae Rich (1808) nom cons

Leoniaceae ADC (1844) = ViolaceaeLeonticaceae Bercht amp J Presl (1820) =

BerberidaceaeLepidobotryaceae JLeacuteonard (1950) nom consLepuropetalaceae Nakai (1943) optional syn-

onym of ParnassiaceaeLilaeaceae Dumort (1829) nom cons =

JuncaginaceaeLiliaceae Juss (1789) nom consLimnanthaceae RBr (1833) nom consLimnocharitaceae Takht ex Cronquist (1981)Limoniaceae Ser (1851) nom cons =

PlumbaginaceaeLinaceae DC ex Perleb (1818) nom consLindenbergiaceae Doweld (2001) = Oroban-

chaceaeLinnaeaceae (Raf) Backlund (1998) optional

synonym of CaprifoliaceaeLiriodendraceae FABarkley (1975) =

MagnoliaceaeLissocarpaceae Gilg (1924) nom cons =

EbenaceaeLoasaceae Juss (1804) nom consLobeliaceae Juss (1813) nom cons optional

synonym of CampanulaceaeLoganiaceae RBr (1814) nom consLomandraceae Lotsy (1911) = Laxmanniaceae

optional synonym of AsparagaceaeLophiolaceae Nakai (1943) = NartheciaceaeLophiraceae Loud (1830) = OchnaceaeLophophytaceae (Schott amp Endl) Bromhead

(1840) = BalanophoraceaeLophopyxidaceae (Engl) HPfeiff (1951)Loranthaceae Juss (1808) nom consLowiaceae Ridl (1924) nom consLuxemburgiaceae Soler (1908) = OchnaceaeLuzuriagaceae Lotsy (1911)Lyginiaceae BGBriggs amp LASJohnson (2000) =

AnarthriaceaeLythraceae JSt-Hil (1805) nom consMackinlayaceae Doweld (2001)Maesaceae (ADC) Anderb BStaringhl amp Kaumlllersjouml

(2000)Magnoliaceae Juss (1789) nom consMalaceae Small (1903) nom cons = RosaceaeMalesherbiaceae DDon (1827) nom cons

optional synonym of PassifloraceaeMalpighiaceae Juss (1789) nom consMalvaceae Juss (1789) nom consMarantaceae RBr (1814) nom consMarcgraviaceae Juss ex DC (1816) nom consMartyniaceae Horan (1847) nom consMastixiaceae Calest (1905) = CornaceaeMaundiaceae Nakai (1943) = JuncaginaceaeMayacaceae Kunth (1842) nom consMedeolaceae (SWatson) Takht (1987) = Liliaceae

432 AGP II


Medusagynaceae Engl amp Gilg (1924) nomcons optional synonym of Ochnaceae

Medusandraceae Brenan (1952) nom consunplaced

Melanophyllaceae Takht ex Airy Shaw (1972)Melanthiaceae Batsch ex Borkh (1796) nom

consMelastomataceae Juss (1789) nom consMeliaceae Juss (1789) nom consMelianthaceae Bercht amp JPresl (1820) nom

consMeliosmaceae Endl (1841) = SabiaceaeMemecylaceae DC (1827) nom cons optional

synonym of MelastomataceaeMendonciaceae Bremek (1954) = AcanthaceaeMenispermaceae Juss (1789) nom consMenyanthaceae Bercht amp JPresl (1820) nom

consMesembryanthemaceae Fenzl (1836) nom cons =

AizoaceaeMetteniusaceae HKarst ex Schnizl (1860-1870)

unplacedMeyeniaceae Sreem (1977) = AcanthaceaeMilulaceae Traub (1972) = AlliaceaeMimosaceae RBr (1814) nom cons = FabaceaeMisodendraceae JAgardh (1858) nom

consMitrastemonaceae Makino (1911) nom cons

unplacedMolluginaceae Bartl (1825) nom consMonimiaceae Juss (1809) nom consMonotaceae Kosterm (1989) = DipterocarpaceaeMonotropaceae Nutt (1818) nom cons =

EricaceaeMontiniaceae Nakai (1943) nom consMoraceae Link (1831) nom consMorinaceae Raf (1820) optional synonym of

CaprifoliaceaeMoringaceae Martynov (1820) nom consMouririaceae Gardner (1840) = Memecylaceae

optional synonym of MelastomataceaeMoutabeaceae Endl (1841) = PolygalaceaeMuntingiaceae CBayer MWChase amp MFFay

(1998)Musaceae Juss (1789) nom consMyodocarpaceae Doweld (2001)Myoporaceae RBr (1810) nom cons =

ScrophulariaceaeMyricaceae ARich ex Kunth (1817) nom

consMyriophyllaceae Schultz Sch (1832) =

HaloragaceaeMyristicaceae RBr (1810) nom consMyrothamnaceae Nied (1891) nom cons

optional synonym of GunneraceaeMyrsinaceae RBr (1810) nom cons

Myrtaceae Juss (1789) nom consMystropetalaceae Hookf (1853) =

BalanophoraceaeNajadaceae Juss (1789) nom cons =

HydrocharitaceaeNandinaceae Horan (1834) = BerberidaceaeNapoleonaceae ARich (1827) = LecythidaceaeNartheciaceae Fr ex Bjurzon (1846)Naucleaceae Wernh (1911) = RubiaceaeNectaropetalaceae (HWinkl) Exell amp Mendonccedila

(1951) = ErythroxylaceaeNelsoniaceae Sreem (1977) = AcanthaceaeNelumbonaceae Bercht amp JPresl (1820) nom

consNemacladaceae Nutt (1842) = Lobeliaceae

optional synonym of CampanulaceaeNepenthaceae Bercht amp JPresl (1820) nom

consNesogenaceae Marais (1981) = OrobanchaceaeNeuradaceae Link (1831) nom consNeuwiediaceae (Burns-Bal amp VAFunk)

RDahlgren ex Reveal amp Hoogland (1991) =Orchidaceae

Nitrariaceae Bercht amp JPresl (1820) nomcons

Nolanaceae Dumort (1829) nom cons =Solanaceae

Nolinaceae Nakai (1943) = Ruscaceae optionalsynonym of Asparagaceae

Nothofagaceae Kuprian (1962)Nupharaceae AKern (1891) = NymphaeaceaeNyctaginaceae Juss (1789) nom consNyctanthaceae JAgardh (1858) = OleaceaeNymphaeaceae Salisb (1805) nom consNypaceae Brongn ex Le Maout amp Decne (1868) =

ArecaceaeNyssaceae Juss ex Dumort (1829) nom cons

optional synonym of CornaceaeOchnaceae DC (1811) nom consOctoknemaceae Soler (1908) nom cons =

OlacaceaeOftiaceae Takht amp Reveal (1993) = Scrophulari-

aceaeOlacaceae RBr (1818) nom consOleaceae Hoffmanns amp Link (1809) nom

consOliniaceae Arn (1839) nom consOnagraceae Juss (1789) nom consOncothecaceae Kobuski ex Airy Shaw (1964)Ophiopogonaceae Endl (1841) = Ruscaceae

optional synonym of AsparagaceaeOpiliaceae Valeton (1886) nom consOrchidaceae Juss (1789) nom consOrobanchaceae Vent (1799) nom consOrontiaceae Bartl (1830) = AraceaeOxalidaceae RBr (1818) nom cons



Oxystylidaceae Hutch (1969) = BrassicaceaePachysandraceae JAgardh (1858) = BuxaceaePaeoniaceae Raf (1815) nom consPaivaeusaceae A Meeuse (1990) =

PicrodendraceaePalmae Juss (1789) nom alt et cons = ArecaceaePandaceae Engl amp Gilg (1912-1913) nom consPandanaceae RBr (1810) nom consPangiaceae Endl (1841) = AchariaceaePapaveraceae Juss (1789) nom consPapilionaceae Giseke (1792) nom alt et cons =

FabaceaeParacryphiaceae Airy Shaw (1964)Parnassiaceae Martynov (1820) nom consParonychiaceae Juss (1815) = CaryophyllaceaeParopsiaceae Dumort (1829) = PassifloraceaePassifloraceae Juss ex Roussel (1806) nom

consPaulowniaceae Nakai (1949)Pedaliaceae RBr (1810) nom consPeganaceae (Engl) Tieghm ex Takht (1987)

optional synonym of NitrariaceaePellicieraceae (Triana amp Planch) LBeauvis ex

Bullock (1959) optional synonym ofTetrameristaceae

Penaeaceae Sweet ex Guill (1828) nom consPennantiaceae JAgardh (1858)Pentadiplandraceae Hutch amp Dalziel (1928)Pentaphragmataceae JAgardh (1858) nom

consPentaphylacaceae Engl (1897) nom consPentastemonaceae Duyfjes (1992) = StemonaceaePenthoraceae Rydb ex Britt (1901) nom

cons optional synonym of HaloragaceaePeperomiaceae ACSm (1981) = PiperaceaePeraceae Klotzsch = EuphorbiaceaePeridiscaceae Kuhlm (1950) nom consPeriplocaceae (Kostel) Schltr (1905) nom cons =

ApocynaceaePeripterygiaceae G King (1895) =

CardiopteridaceaePetermanniaceae Hutch (1934) nom cons =

ColchicaceaePetiveriaceae CAgardh (1824) = PhytolaccaceaePetrosaviaceae Hutch (1934) nom consPhellinaceae (Loes) Takht (1967)Philadelphaceae Martynov (1820) =

HydrangeaceaePhilesiaceae Dumort (1829) nom consPhilydraceae Link (1821) nom consPhormiaceae JAgardh (1858) = Hemerocalli-

daceae optional synonym of XanthorrhoeaceaePhrymaceae Schauer (1847) nom consPhyllanthaceae Martynov (1820)Phyllonomaceae Small (1905)Physenaceae Takht (1985)

Phytolaccaceae RBr (1818) nom consPicramniaceae Fernando amp Quinn (1995)Picrodendraceae Small (1917) nom consPiperaceae Bercht amp J Presl (1820) nom consPistiaceae Rich ex CAgardh (1822) = AraceaePittosporaceae RBr (1814) nom consPlagiopteraceae Airy Shaw (1964) = CelastraceaePlantaginaceae Juss (1789) nom consPlatanaceae TLestib (1826) nom cons

optional synonym of ProteaceaePlatycaryaceae Nakai ex Doweld (2001) =

JuglandaceaePlatyspermataceae Doweld (2001) = Escalloni-

aceaePlatystemonaceae (Spach) Lilja (1870) =

PapaveraceaePlocospermataceae Hutch (1973)Plumbaginaceae Juss (1789) nom consPlumeriaceae Horan (1834) = ApocynaceaePoaceae (RBr) Barnh (1895) nom consPodoaceae Baill ex Franch (1889) =

AnacardiaceaePodophyllaceae DC (1817) nom cons =

BerberidaceaePodostemaceae Kunth (1816) nom consPolemoniaceae Juss (1789) nom consPoliothyrsidaceae (GSFan) Doweld (2001) =

SalicaceaePolpodaceae Nakai (1942) = MolluginaceaePolygalaceae Hoffmanns amp Link (1809) nom

consPolygonaceae Juss (1789) nom consPolygonanthaceae Croizat (1943) =

AnisophylleaceaePolyosmaceae Blume (1851)Pontederiaceae Kunth (1816) nom consPorantheraceae (Pax) Hurus (1954) =

PhyllanthaceaePortulacaceae Juss (1789) nom consPortulacariaceae (Fenzl) Doweld (2001) =

PortulacaceaePosidoniaceae Hutch (1934) nom consPotaliaceae Mart (1827) = GentianaceaePotamogetonaceae Rchb (1828) nom consPottingeriaceae (Engl) Takht (1987) unplacedPrimulaceae Batsch ex Borkh (1797) nom consPrioniaceae SLMunro amp HPLinder (1998) =

ThurniaceaePrionotaceae Hutch (1969) = EricaceaeProteaceae Juss (1789) nom consPseudanthaceae Endl (1839) = PicrodendraceaePsiloxylaceae Croizat (1960)Ptaeroxylaceae J-FLeroy (1960) = RutaceaePteridophyllaceae (Murb) Nakai ex Reveal amp

Hoogland (1991) optional synonym ofPapaveraceae

434 AGP II


Pterostemonaceae Small (1905) nom consoptional synonym of Iteaceae

Punicaceae Horan (1834) nom cons =Lythraceae

Putranjivaceae Endl (1841)Pyrolaceae Lindl (1829) nom cons = EricaceaeQuiinaceae Choisy ex Engl (1888) nom cons

optional synonym of OchnaceaeQuillajaceae DDon (1831)Quintiniaceae Doweld (2001) = Sphenostemo-

naceaeRafflesiaceae Dumort (1829) nom cons

unplacedRanunculaceae Juss (1789) nom consRanzaniaceae (Kumaz amp Terab) Takht (1994) =

BerberidaceaeRapateaceae Dumort (1829) nom consReaumuriaceae Ehrenb ex Lindl (1830) =

TamaricaceaeResedaceae Bercht amp JPresl (1820) nom consRestionaceae RBr (1810) nom consRetziaceae Bartl (1830) = StilbaceaeRhabdodendraceae Prance (1968)Rhamnaceae Juss (1789) nom consRhinanthaceae Vent (1799) = OrobanchaceaeRhipogonaceae Conran amp Clifford (1985)Rhizophoraceae Pers (1807) nom cons

optional synonym of ErythroxylaceaeRhodoleiaceae Nakai (1943) = HamamelidaceaeRhoipteleaceae Hand-Mazz (1932) nom cons

optional synonym of JuglandaceaeRhopalocarpaceae Hemsl ex Takht (1987) =

SphaerosepalaceaeRhynchocalycaceae LASJohnson amp

BGBriggs (1985)Rhynchothecaceae Endl (1841) = LedocarpaceaeRoridulaceae Bercht amp JPresl (1820) nom consRosaceae Juss (1789) nom consRousseaceae DC (1839)Roxburghiaceae Wall (1832) = StemonaceaeRubiaceae Juss (1789) nom consRuppiaceae Horan (1834) nom consRuscaceae Spreng (1826) nom cons optional

synonym of AsparagaceaeRutaceae Juss (1789) nom consSabiaceae Blume (1851) nom consSaccifoliaceae Maguire amp Pires (1978) =

GentianaceaeSalazariaceae FABarkley (1975) = LamiaceaeSalicaceae Mirb (1815) nom consSalicorniaceae Martynov (1820) = AmaranthaceaeSalpiglossidaceae Hutch (1969) = SolanaceaeSalsolaceae Menge (1839) = AmaranthaceaeSalvadoraceae Lindl (1836) nom consSambucaceae Batsch ex Borkh (1797) =

Adoxaceae

Samolaceae Raf (1820) = TheophrastaceaeSamydaceae Vent (1799) = SalicaceaeSaniculaceae (Burnett) ALoumlve amp DLoumlve (1974) =

ApiaceaeSansevieriaceae Nakai (1936) = Ruscaceae

optional synonym of AsparagaceaeSantalaceae RBr (1810) nom consSapindaceae Juss (1789) nom consSapotaceae Juss (1789) nom consSarcobataceae Behnke (1997)Sarcolaenaceae Caruel (1881) nom consSarcophytaceae AKern (1891) =

BalanophoraceaeSarcospermataceae HJLam (1925) nom cons =

SapotaceaeSargentodoxaceae Stapf ex Hutch (1926) nom

cons = LardizabalaceaeSarraceniaceae Dumort (1829) nom consSaurauiaceae Griseb (1854) nom cons =

ActinidiaceaeSaururaceae Martynov (1820) nom consSauvagesiaceae Dumort (1829) = OchnaceaeSaxifragaceae Juss (1789) nom consScaevolaceae Lindl (1830) = GoodeniaceaeScepaceae Lindl (1836) = PhyllanthaceaeScheuchzeriaceae FRudolphi (1830) nom

consSchisandraceae Blume (1830) nom consSchlegeliaceae (AHGentry) Reveal (1996)Sclerophylacaceae Miers (1848) = SolanaceaeScoliopaceae Takht (1996) = LiliaceaeScrophulariaceae Juss (1789) nom consScybaliaceae AKern (1891) = BalanophoraceaeScyphostegiaceae Hutch (1926) nom cons =

SalicaceaeScytopetalaceae Engl (1897) nom cons =

LecythidaceaeSelaginaceae Choisy (1823) nom cons =

ScrophulariaceaeSesamaceae RBr ex Bercht amp JPresl (1820) =

PedaliaceaeSesuviaceae Horan (1834) = AizoaceaeSetchellanthaceae Iltis (1999)Simaroubaceae DC (1811) nom consSimmondsiaceae Tiegh (1899)Sinofranchetiaceae Doweld (2001) = Lardiza-

balaceaeSiparunaceae (ADC) Schodde (1970)Siphonodontaceae (Croizat) Gagnep amp Tardieu

ex Tardieu (1951) nom cons = CelastraceaeSladeniaceae Airy Shaw (1964) optional

synonym of PentaphylacaceaeSmilacaceae Vent (1799) nom consSolanaceae Juss (1789) nom consSonneratiaceae Engl (1897) nom cons =

Lythraceae



Sparganiaceae Hanin (1811) nom consSpergulaceae Bartl (1825) = CaryophyllaceaeSphaerosepalaceae (Warb) Tiegh ex Bullock

(1959)Sphenocleaceae (Lindl) Baskerville (1839)

nom consSphenostemonaceae PRoyen amp Airy Shaw

(1972)Spigeliaceae Mart (1827) = LoganiaceaeSpiraeaceae Bertuch (1801) = RosaceaeStachyuraceae JAgardh (1858) nom consStackhousiaceae RBr (1814) nom cons =

CelastraceaeStaphyleaceae Martynov (1820) nom consStaticaceae Cassel (1817) = PlumbaginaceaeStegnospermataceae Nakai (1942)Stemonaceae Caruel (1878) nom consStemonuraceae (MRoem) Karingrehed (2001)Stenomeridaceae JAgardh (1858) =

DioscoreaceaeSterculiaceae Vent ex Salisb (1807) nom cons =

MalvaceaeStilaginaceae CAgardh (1824) = EuphorbiaceaeStilbaceae Kunth (1831) nom consStrasburgeriaceae Soler (1908) nom consStrelitziaceae Hutch (1934) nom consStreptochaetaceae Nakai (1943) = PoaceaeStrychnaceae DC ex Perleb (1818) = LoganiaceaeStylidiaceae RBr (1810) nom consStylobasiaceae JAgardh (1858) = SurianaceaeStylocerataceae (Pax) Takht ex Reveal amp

Hoogland (1990) = BuxaceaeStyracaceae DC amp Spreng (1821) nom

consSurianaceae Arn (1834) nom consSymphoremataceae (Meisn) Mold ex Reveal amp

Hoogland (1991) = LamiaceaeSymplocaceae Desf (1820) nom consTaccaceae Bercht amp JPresl (1820) nom cons =

DioscoreaceaeTakhtajaniaceae (J-FLeroy) J-FLeroy (1980) =

WinteraceaeTalinaceae (Fenzl) Doweld (2001) = PortulacaceaeTamaricaceae Bercht amp JPresl (1820) nom

consTapisciaceae (Pax) Takht (1987)Tecophilaeaceae Leyb (1862) nom consTepuianthaceae Maguire amp Steyerm (1981) =

ThymelaeaceaeTernstroemiaceae Mirb ex DC (1816)

optional synonym of PentaphylacaceaeTetracarpaeaceae Nakai (1943) optional syn-

onym of HaloragaceaeTetracentraceae ACSm (1945) nom cons

optional synonym of TrochodendraceaeTetrachondraceae Wettst (1924)

Tetradiclidaceae (Engl) Takht (1986) anoptional synonym of Nitrariaceae

Tetragoniaceae Link (1831) nom cons =Aizoaceae

Tetramelaceae Airy Shaw (1964)Tetrameristaceae Hutch (1959)Tetrastylidiaceae Tiegh (1899) = OlacaceaeThalassiaceae Nakai (1943) = HydrocharitaceaeThalictraceae Raf (1815) = RanunculaceaeTheaceae Mirb ex Ker Gawl (1816) nom consTheligonaceae Dumort (1829) nom cons =

RubiaceaeThemidaceae Salisb (1866) optional synonym

of AsparagaceaeTheophrastaceae Link (1829) nom consThismiaceae JAgardh (1858) nom cons =

BurmanniaceaeThomandersiaceae Sreem (1977) = AcanthaceaeThunbergiaceae (Dumort) Lilja (1870) =

AcanthaceaeThurniaceae Engl (1907) nom consThymelaeaceae Juss (1789) nom consTicodendraceae Goacutemez-Laur amp LDGoacutemez

(1991)Tiliaceae Juss (1789) nom cons = MalvaceaeTofieldiaceae Takht (1995)Torricelliaceae Hu (1934)Tovariaceae Pax (1891) nom consTrapaceae Dumort (1829) nom cons =

LythraceaeTrapellaceae Honda amp Sakis (1930) = PedaliaceaeTremandraceae RBr ex DC (1824) nom cons =

ElaeocarpaceaeTrewiaceae Lindl (1836) = EuphorbiaceaeTribelaceae Airy Shaw (1964)Tribulaceae Trautv (1853) = ZygophyllaceaeTrichopodaceae Hutch (1934) nom cons =

DioscoreaceaeTricyrtidaceae Takht (1997) nom cons =

LiliaceaeTrigoniaceae Endl (1841) nom cons optional

synonym of ChrysobalanaceaeTrilliaceae Chevall (1827) nom cons =

MelanthiaceaeTrimeniaceae LSGibbs (1917) nom consTriplostegiaceae AE Bobrov ex Airy Shaw (1964)

= Dipsaceaceae optional synonym ofCaprifoliaceae

Tristichaceae Willis (1915) = PodostemaceaeTriuridaceae Gardner (1843) nom consTrochodendraceae Eichler (1865) nom consTropaeolaceae Bercht amp JPresl (1820) nom

consTulbaghiaceae Salisb (1866) = AlliaceaeTurneraceae Kunth ex DC (1828) nom cons

optional synonym of Passifloraceae

436 AGP II


Typhaceae Juss (1789) nom consUapacaceae Airy Shaw (1964) = PhyllanthaceaeUlmaceae Mirb (1815) nom consUmbelliferae Juss (1789) nom alt et cons =

ApiaceaeUrticaceae Juss (1789) nom consUvulariaceae AGray ex Kunth (1843) nom cons

= ColchicaceaeVacciniaceae DC ex Perleb (1818) nom cons =

EricaceaeVahliaceae Dandy (1959)Valerianaceae Batsch (1802) nom cons

optional synonym of CaprifoliaceaeVallisneriaceae Link (1829) = HydrocharitaceaeVelloziaceae Hook (1827) nom consVerbascaceae Raf (1821) = ScrophulariaceaeVerbenaceae JSt-Hil (1805) nom consVeronicaceae Cassel (1817) = PlantaginaceaeViburnaceae Raf (1820) = AdoxaceaeViolaceae Batsch (1802) nom consViscaceae Batsch (1802) = SantalaceaeVitaceae Juss (1789) nom cons

Viticaceae Juss (1789) = LamiaceaeVivianiaceae Klotzsch (1836)Vochysiaceae ASt-Hil (1820) nom consWalleriaceae (RDahlgren) Takht (1995) nom

cons = TecophilaeaceaeWellstediaceae (Pilg) Novaacutek (1943) =

BoraginaceaeWinteraceae RBr ex Lindl (1830) nom

consXanthophyllaceae (Baill) Gagnep ex Reveal amp

Hoogland (1990) = PolygalaceaeXanthorrhoeaceae Dumort (1829) nom

consXeronemataceae MWChase Rudall amp MFFay

(2001)Xerophyllaceae Takht (1996) = MelanthiaceaeXyridaceae CAgardh (1823) nom consZannichelliaceae Chevall (1827) nom cons =

PotamogetonaceaeZingiberaceae Martynov (1820) nom consZosteraceae Dumort (1829) nom consZygophyllaceae RBr (1814) nom cons

400

AGP II



2003

141

399ndash436

development of phylogenetic theory and application toanalysis of large data sets (eg Hillis 1996) and var-ious methods for estimating the support for individualclades in the phylogenetic trees (Felsenstein 1985Farris

et al

1996) The outline of a phylogenetic treeof all flowering plants became established and severalwell supported major clades involving many familiesof flowering plants were identified In many cases thenew knowledge of phylogeny revealed relationships inconflict with the then widely used modern classifica-tions (eg Cronquist 1981 Thorne 1992 Takhtajan1997) which were based on selected similarities anddifferences in morphology rather than cladistic anal-ysis of larger data sets involving DNA sequences orother forms of systematic data It became clear thatnone of the previous classifications accuratelyreflected phylogenetic relationships of floweringplants and communication about plant evolutionreferring to the old classification schemes becameincreasingly difficult To alleviate this problem agroup of flowering plant systematists calling them-selves the Angiosperm Phylogeny Group (APG forshort) proposed a new classification for the families offlowering plants (APG 1998)

The initial APG (1998) system comprised 462 fami-lies arranged in 40 putatively monophyletic ordersand a few monophyletic higher groups The latter werenamed informally as monocots commelinoids (herechanged to commelinids to prevent confusion withsubfamily Commelinoideae of Commelinaceae) eud-icots core eudicots rosids including eurosids I and IIand asterids including euasterids I and II The focuswas on orders and less on families An attempt wasmade to recognize orders well supported as monophyl-etic in large jackknife analyses of molecular data(Kaumlllersjouml

et al

1998) In general the orders werefairly widely circumscribed especially in comparisonwith those of Takhtajan (1997) A few monofamilialorders were recognized (Ceratophyllales Acorales andArecales) for cases in which these families were appar-ently sister groups of larger clades including severalorders Many families were not classified to orderbecause their positions were uncertain or unknownand these families were listed under the supraordinalgroups where they were known to belong or at the endof the system in a list of families probably eudicots ofuncertain position APG predicted that there would belittle need to change the circumscription of the ordersexcept for inclusion of families not then assigned toorder and possible transfer of occasional misplacedfamilies It was also realized that new orders might beestablished if monophyletic groups of unplaced fami-lies were identified

The APG system also involved the recognition ofstrictly monophyletic groups at all levels but it wasacknowledged that there were families known to be

non-monophyletic (eg Euphorbiaceae and Scrophu-lariaceae) Reclassification of these into monophyleticunits was not possible in 1998 and required furtherinvestigation Furthermore monophyly of many fam-ilies remained to be investigated with extensive sam-pling and application of molecular phylogenetictechniques Thus it was acknowledged that somechanges in family circumscription would be necessaryto reflect improved understanding of phylogeneticrelationships For some families already investigatedand found to be monophyletic alternative optionalcircumscriptions were indicated by listing the sisterfamily or families in square brackets immediatelyafter the family For example Nymphaeaceae could beinterpreted either to exclude or include a sister familyCabombaceae

Five years have now passed since the APG systemwas compiled Recent advances in our knowledge offlowering plant phylogeny indeed have motivated sev-eral changes in family circumscription and classifica-tion as well as the addition of a few new orders Wetherefore present here an updated version of the APGsystem

In general we have adopted a conservativeapproach and propose here changes in the APG sys-tem only when there is substantial new evidence sup-porting a revised classification Five additional ordersare recognized Austrobaileyales Canellales Celas-trales Crossosomatales and Gunnerales These rep-resent well-supported monophyletic groups offamilies unclassified to order in APG (1998) Circum-scription of none of the APG orders has been changedexcept for the addition of a number of the familiesunclassified to order in APG (1998) When morerecent analyses have demonstrated that such familiesof formerly uncertain position are well nested insidethe APG orders or well supported as sister groups toany of the APG orders the latter have been expandedto include these families Thus some APG ordershave been more widely circumscribed to include theirsister groups (eg Adoxaceae being included in Dipsa-cales cf Bremer 2000) except in one case in whichthe pair of Canellaceae and Winteraceae has beenestablished as an order Canellales rather thanincluded in their sister group Piperales (mostresearchers would consider these two groups toodivergent to include in a single order) No APG ordershave been merged or split and no families have beentransferred from one order to another Only in onecase has a family been removed from an APG orderOncothecaceae have been excluded from Garryalesand assigned to a position at the beginning of theeuasterids I without classification to order becauserecent analyses have not supported any clear (iebootstrap- or jackknife-supported) ordinal position forthat family


401



2003

141

399ndash436

Figure 1


et al


rbcL

and

atpB


+

PiperalesLaurales

+


et al


rbcL


et al


et al


402

AGP II



2003

141

399ndash436


rbcL

and

atpB


et al


et al


et al

2002)and of

rbcL


et al


et al


et al

2000 Soltis

et al




et al


et al




Amborella


et al

1999 Soltis

et al


et al


Amborella


et al


Amborella


Amborella


Austrobaileya

Illicium

and

Schisandra


rbcL

atpB


et al

2000b) Material of

Trimenia


rbcL


et al


rbcL

atpB


et al


Trimeni


et al

1999 Zanis

et al



et al

(1999) and Soltis

et al


et al

2003) and 67 (Qiu

et al


et al


et al



403



2003

141

399ndash436


et al

1999 100 Zanis

et al


et al

1999 100 Zanis

et al


+



et al

1999 Qiu

et al

1999 Soltis

et al

1999 Zanis

et al


lt


et al


et al


Lactoris


Aristolochia

+

Thottea

(Qiu

et al

1999 Zanis

et al

2003) or

Aristolochia

alone (Soltis

et al

2000a)

Thottea


Lactoris


Lactoris


Lactoris



+


et al


MONOCOTS


et al



et al

(2000) and Soltis

et al


matK


Japanolirion


et al


rbcL atpB




404 AGP II










EUDICOTS








406 AGP II





ROSIDS

















408 AGP II





ASTERIDS














410 AGP II













CONCLUSION



REFERENCES










412 AGP II

































































414 AGP II


























































APPENDIX



416 AGP II





cons]




MAGNOLIIDS








MONOCOTS










(2001)





COMMELINIDS







EUDICOTS







CORE EUDICOTS



418 AGP II









ROSIDS







EUROSIDS I



















EUROSIDS II




cons


420 AGP II







ASTERIDS








EUASTERIDS I










cons

EUASTERIDS II






















422 AGP II


































Ranunculales












424 AGP II











































426 AGP II







under eurosid I
























































428 AGP II





































































430 AGP II













































































432 AGP II





































































434 AGP II
















Adoxaceae
















Lythraceae






































436 AGP II
















401



2003

141

399ndash436

Figure 1


et al


rbcL

and

atpB


+

PiperalesLaurales

+


et al


rbcL


et al


et al


402

AGP II



2003

141

399ndash436


rbcL

and

atpB


et al


et al


et al

2002)and of

rbcL


et al


et al


et al

2000 Soltis

et al




et al


et al




Amborella


et al

1999 Soltis

et al


et al


Amborella


et al


Amborella


Amborella


Austrobaileya

Illicium

and

Schisandra


rbcL

atpB


et al

2000b) Material of

Trimenia


rbcL


et al


rbcL

atpB


et al


Trimeni


et al

1999 Zanis

et al



et al

(1999) and Soltis

et al


et al

2003) and 67 (Qiu

et al


et al


et al



403



2003

141

399ndash436


et al

1999 100 Zanis

et al


et al

1999 100 Zanis

et al


+



et al

1999 Qiu

et al

1999 Soltis

et al

1999 Zanis

et al


lt


et al


et al


Lactoris


Aristolochia

+

Thottea

(Qiu

et al

1999 Zanis

et al

2003) or

Aristolochia

alone (Soltis

et al

2000a)

Thottea


Lactoris


Lactoris


Lactoris



+


et al


MONOCOTS


et al



et al

(2000) and Soltis

et al


matK


Japanolirion


et al


rbcL atpB




404 AGP II










EUDICOTS








406 AGP II





ROSIDS

















408 AGP II





ASTERIDS














410 AGP II













CONCLUSION



REFERENCES










412 AGP II

































































414 AGP II


























































APPENDIX



416 AGP II





cons]




MAGNOLIIDS








MONOCOTS










(2001)





COMMELINIDS







EUDICOTS







CORE EUDICOTS



418 AGP II









ROSIDS







EUROSIDS I



















EUROSIDS II




cons


420 AGP II







ASTERIDS








EUASTERIDS I










cons

EUASTERIDS II






















422 AGP II


































Ranunculales












424 AGP II











































426 AGP II







under eurosid I
























































428 AGP II





































































430 AGP II













































































432 AGP II





































































434 AGP II
















Adoxaceae
















Lythraceae






































436 AGP II















402

AGP II



2003

141

399ndash436


rbcL

and

atpB


et al


et al


et al

2002)and of

rbcL


et al


et al


et al

2000 Soltis

et al




et al


et al




Amborella


et al

1999 Soltis

et al


et al


Amborella


et al


Amborella


Amborella


Austrobaileya

Illicium

and

Schisandra


rbcL

atpB


et al

2000b) Material of

Trimenia


rbcL


et al


rbcL

atpB


et al


Trimeni


et al

1999 Zanis

et al



et al

(1999) and Soltis

et al


et al

2003) and 67 (Qiu

et al


et al


et al



403



2003

141

399ndash436


et al

1999 100 Zanis

et al


et al

1999 100 Zanis

et al


+



et al

1999 Qiu

et al

1999 Soltis

et al

1999 Zanis

et al


lt


et al


et al


Lactoris


Aristolochia

+

Thottea

(Qiu

et al

1999 Zanis

et al

2003) or

Aristolochia

alone (Soltis

et al

2000a)

Thottea


Lactoris


Lactoris


Lactoris



+


et al


MONOCOTS


et al



et al

(2000) and Soltis

et al


matK


Japanolirion


et al


rbcL atpB




404 AGP II










EUDICOTS








406 AGP II





ROSIDS

















408 AGP II





ASTERIDS














410 AGP II













CONCLUSION



REFERENCES










412 AGP II

































































414 AGP II


























































APPENDIX



416 AGP II





cons]




MAGNOLIIDS








MONOCOTS










(2001)





COMMELINIDS







EUDICOTS







CORE EUDICOTS



418 AGP II









ROSIDS







EUROSIDS I



















EUROSIDS II




cons


420 AGP II







ASTERIDS








EUASTERIDS I










cons

EUASTERIDS II






















422 AGP II


































Ranunculales












424 AGP II











































426 AGP II







under eurosid I
























































428 AGP II





































































430 AGP II













































































432 AGP II





































































434 AGP II
















Adoxaceae
















Lythraceae






































436 AGP II
















403



2003

141

399ndash436


et al

1999 100 Zanis

et al


et al

1999 100 Zanis

et al


+



et al

1999 Qiu

et al

1999 Soltis

et al

1999 Zanis

et al


lt


et al


et al


Lactoris


Aristolochia

+

Thottea

(Qiu

et al

1999 Zanis

et al

2003) or

Aristolochia

alone (Soltis

et al

2000a)

Thottea


Lactoris


Lactoris


Lactoris



+


et al


MONOCOTS


et al



et al

(2000) and Soltis

et al


matK


Japanolirion


et al


rbcL atpB




404 AGP II










EUDICOTS








406 AGP II





ROSIDS

















408 AGP II





ASTERIDS














410 AGP II













CONCLUSION



REFERENCES










412 AGP II

































































414 AGP II


























































APPENDIX



416 AGP II





cons]




MAGNOLIIDS








MONOCOTS










(2001)





COMMELINIDS







EUDICOTS







CORE EUDICOTS



418 AGP II









ROSIDS







EUROSIDS I



















EUROSIDS II




cons


420 AGP II







ASTERIDS








EUASTERIDS I










cons

EUASTERIDS II






















422 AGP II


































Ranunculales












424 AGP II











































426 AGP II







under eurosid I
























































428 AGP II





































































430 AGP II













































































432 AGP II





































































434 AGP II
















Adoxaceae
















Lythraceae






































436 AGP II















404 AGP II










EUDICOTS








406 AGP II





ROSIDS

















408 AGP II





ASTERIDS














410 AGP II













CONCLUSION



REFERENCES










412 AGP II

































































414 AGP II


























































APPENDIX



416 AGP II





cons]




MAGNOLIIDS








MONOCOTS










(2001)





COMMELINIDS







EUDICOTS







CORE EUDICOTS



418 AGP II









ROSIDS







EUROSIDS I



















EUROSIDS II




cons


420 AGP II







ASTERIDS








EUASTERIDS I










cons

EUASTERIDS II






















422 AGP II


































Ranunculales












424 AGP II











































426 AGP II







under eurosid I
























































428 AGP II





































































430 AGP II













































































432 AGP II





































































434 AGP II
















Adoxaceae
















Lythraceae






































436 AGP II


















EUDICOTS








406 AGP II





ROSIDS

















408 AGP II





ASTERIDS














410 AGP II













CONCLUSION



REFERENCES










412 AGP II

































































414 AGP II


























































APPENDIX



416 AGP II





cons]




MAGNOLIIDS








MONOCOTS










(2001)





COMMELINIDS







EUDICOTS







CORE EUDICOTS



418 AGP II









ROSIDS







EUROSIDS I



















EUROSIDS II




cons


420 AGP II







ASTERIDS








EUASTERIDS I










cons

EUASTERIDS II






















422 AGP II


































Ranunculales












424 AGP II











































426 AGP II







under eurosid I
























































428 AGP II





































































430 AGP II













































































432 AGP II





































































434 AGP II
















Adoxaceae
















Lythraceae






































436 AGP II















406 AGP II





ROSIDS

















408 AGP II





ASTERIDS














410 AGP II













CONCLUSION



REFERENCES










412 AGP II

































































414 AGP II


























































APPENDIX



416 AGP II





cons]




MAGNOLIIDS








MONOCOTS










(2001)





COMMELINIDS







EUDICOTS







CORE EUDICOTS



418 AGP II









ROSIDS







EUROSIDS I



















EUROSIDS II




cons


420 AGP II







ASTERIDS








EUASTERIDS I










cons

EUASTERIDS II






















422 AGP II


































Ranunculales












424 AGP II











































426 AGP II







under eurosid I
























































428 AGP II





































































430 AGP II













































































432 AGP II





































































434 AGP II
















Adoxaceae
















Lythraceae






































436 AGP II

























408 AGP II





ASTERIDS














410 AGP II













CONCLUSION



REFERENCES










412 AGP II

































































414 AGP II


























































APPENDIX



416 AGP II





cons]




MAGNOLIIDS








MONOCOTS










(2001)





COMMELINIDS







EUDICOTS







CORE EUDICOTS



418 AGP II









ROSIDS







EUROSIDS I



















EUROSIDS II




cons


420 AGP II







ASTERIDS








EUASTERIDS I










cons

EUASTERIDS II






















422 AGP II


































Ranunculales












424 AGP II











































426 AGP II







under eurosid I
























































428 AGP II





































































430 AGP II













































































432 AGP II





































































434 AGP II
















Adoxaceae
















Lythraceae






































436 AGP II
























410 AGP II













CONCLUSION



REFERENCES










412 AGP II

































































414 AGP II


























































APPENDIX



416 AGP II





cons]




MAGNOLIIDS








MONOCOTS










(2001)





COMMELINIDS







EUDICOTS







CORE EUDICOTS



418 AGP II









ROSIDS







EUROSIDS I



















EUROSIDS II




cons


420 AGP II







ASTERIDS








EUASTERIDS I










cons

EUASTERIDS II






















422 AGP II


































Ranunculales












424 AGP II











































426 AGP II







under eurosid I
























































428 AGP II





































































430 AGP II













































































432 AGP II





































































434 AGP II
















Adoxaceae
















Lythraceae






































436 AGP II



















CONCLUSION



REFERENCES










412 AGP II

































































414 AGP II


























































APPENDIX



416 AGP II





cons]




MAGNOLIIDS








MONOCOTS










(2001)





COMMELINIDS







EUDICOTS







CORE EUDICOTS



418 AGP II









ROSIDS







EUROSIDS I



















EUROSIDS II




cons


420 AGP II







ASTERIDS








EUASTERIDS I










cons

EUASTERIDS II






















422 AGP II


































Ranunculales












424 AGP II











































426 AGP II







under eurosid I
























































428 AGP II





































































430 AGP II













































































432 AGP II





































































434 AGP II
















Adoxaceae
















Lythraceae






































436 AGP II















412 AGP II

































































414 AGP II


























































APPENDIX



416 AGP II





cons]




MAGNOLIIDS








MONOCOTS










(2001)





COMMELINIDS







EUDICOTS







CORE EUDICOTS



418 AGP II









ROSIDS







EUROSIDS I



















EUROSIDS II




cons


420 AGP II







ASTERIDS








EUASTERIDS I










cons

EUASTERIDS II






















422 AGP II


































Ranunculales












424 AGP II











































426 AGP II







under eurosid I
























































428 AGP II





































































430 AGP II













































































432 AGP II





































































434 AGP II
















Adoxaceae
















Lythraceae






































436 AGP II
















































414 AGP II


























































APPENDIX



416 AGP II





cons]




MAGNOLIIDS








MONOCOTS










(2001)





COMMELINIDS







EUDICOTS







CORE EUDICOTS



418 AGP II









ROSIDS







EUROSIDS I



















EUROSIDS II




cons


420 AGP II







ASTERIDS








EUASTERIDS I










cons

EUASTERIDS II






















422 AGP II


































Ranunculales












424 AGP II











































426 AGP II







under eurosid I
























































428 AGP II





































































430 AGP II













































































432 AGP II





































































434 AGP II
















Adoxaceae
















Lythraceae






































436 AGP II















an update of the angiosperm phylogeny group classiï¬cation for

Documents