(APG, 1998) An Ordinal Classification For The Families of Flowering Plants (APG I)

An Ordinal Classification for the Families of Flowering Plants
Author(s): The Angiosperm Phylogeny Group

Source: Annals of the Missouri Botanical Garden, Vol. 85, No. 4 (1998), pp. 531-553
Published by: Missouri Botanical Garden Press
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Volume85
Number4
1998
Annals
of the
Missouri
Botanical
Garden
AN ORDINAL
CLASSIFICATION FOR THE
FAMILIES OF FLOWERING
PLANTS
The AngiospermPhylogeny Group'
ABSTRACT
Recentcladisticanalysesare revealingthe phylogeny
of flowering
plantsin increasingdetail,and thereis support
forthe monophyly
ofmanymajorgroupsabove thefamilylevel. Withmanyelementsofthemajorbranching
sequence
ofphylogeny
established,a revisedsuprafamilial
offlowering
classification
plantsbecomesbothfeasibleand desirable.
Here we presenta classificationof 462 flowering
plantfamiliesin 40 putativelymonophyletic
ordersand a small
numberof monophyletic,
informal
highergroups.The latterare the monocots,commelinoids,
eudicots,core eudicots,
rosidsincludingeurosidsI and II, and asteridsincludingeuasteridsI and II. Underthese informal
groupsthereare
also listeda numberoffamilieswithoutassignment
to order.At the end ofthe systemis an additionallistoffamilies
of uncertainpositionforwhichno firmdata existregarding
placementanywhere
withinthe system.
Why rearrangefamilies,still less formalizeorders? Higher-levelclassifications,

the groupingof
species into families,orders,etc., are needed as
referencetoolsnot onlyin systematics
but also in
manyotherbranchesofbiology.Knowledgeofphylogeneticrelationshipsof major groupsof organisms,thatis, a phylogenetic
perspective,
is becoming increasingly
important,
and hence theneed for
a phylogenetic
classificationas a referencetool is
also becomingimperative.
Our primary
focusis on orderswitha secondary
emphasison familiesofflowering
plants.The family
is centralin flowering
plant systematics.For example, in studyingan unknownplant we usually
firstidentify
it to family.The orders,on the other
hand, have untilquite recentlybeen of littleimportance,eitherbeingmorphologically
unrecognizable or in mostcases lackingany evolutionary
coherence (Heywood, 1977; MerxmUller,1977).
However,ordersare usefulin teaching,forstudying
1 Recommended
citation,abbreviatedas "APG, 1998." This paper was compiledby KdreBremer,MarkW. Chase,
and PeterF. Stevens,equally responsibleand listedherein alphabeticalorderonly,withcontributions
fromArneA.
Anderberg,
AndersBacklund,BirgittaBremer,BarbaraG. Briggs,PeterK. Endress,MichaelF. Fay,PeterGoldblatt,
Mats H. G. Gustafsson,
Sara B. Hoot,WalterS. Judd,MariKdllersj6,ElizabethA. Kellogg,KathleenA. Kron,Donald
H. Les, CynthiaM. Morton,Daniel L. Nickrent,RichardG. Olmstead,RobertA. Price,Christopher
J. Quinn,James
E. Rodman,Paula J. Rudall, VincentSavolainen,Douglas E. Soltis,Pamela S. Soltis,KennethJ. Sytsma,and Mats
Thulin(in alphabeticalorder).Addresses:K. Bremer,Department
ofSystematic
Botany,Uppsala University,
Villavagen
6, S-752 36 Uppsala,Sweden;M. W. Chase, JodrellLaboratory,
RoyalBotanicGardens,Kew,Richmond,Surrey,
TW9
3DS, U.K.; P. F. Stevens,HarvardUniversity
Herbaria,22 DivinityAvenue,Cambridge,Massachusetts02138, U.S.A.
ANN. MISSOURI BOT. GARD. 85: 531-553.
1998.
532
Annals of the
MissouriBotanical Garden
We note thatthe selectionof clades to be repand in positioninggeneraof

familyrelationships,
from
is different
The didacticvalue of suprafami- resentedin a formalclassification
doubtfulaffinity.
has been emphasizedby variousau- the procedureof namingthese clades. The latter
lial groupings
is
in phylogenetics
thors(e.g., Dahlgren,1975; Thorne,1976; Davis, issue ofbiologicalnomenclature
muchdebated(e.g., Cantinoet al., 1997;
1978; Takhtajan,1997). This value is even more currently
offlowering
plants de Queiroz,1997; Lid6n et al., 1997), butwe have
evidentnowthatthe phylogeny
naming"sensu de
is beingdisclosedin increasingdetail.Manyofthe not adoptedany "phylogenetic
ordersrecognizedby earlierauthorsare notmono- Queirozand Gauthier(1994). We operateunderthe
Code ofBotanicalNomenclaphyletic,yetthereis a pressingneed fornamesto currentInternational
communicate the knowledge of monophyletic ture(Greuteret al., 1994) and chooseto emphasize
groupingsof familiesthat are becomingevident. the ranksoffamilyand order.The LinnaeancateWith the major branchingsequence of flowering goriesserveas a convenientmnemonicdevice for
becomingclearer,a revisedfamil- rememberinghierarchical relationships,but it
plantphylogeny
shouldofcoursebe realizedthatgroupsofthesame
is feasible.
ial and ordinalclassification
unitsunnon-comparable
Floweringplant classificationsystemsfromthe rank are evolutionarily
late 1970s seemed to be stable and showsubstan- less theyare sistergroups.
problemswhenestablishThere are noteworthy
tial agreement,but this stabilityhas been rudely
shatteredas newkindsofdata and newmethodsof ing the namesfortaxa at ordinaland otherhigher
has
littleattention
es- taxonomiclevels. Untilrecently,
data have becomefirmly
analyzingconventional
at theselevels,and
tablished(Stevens,1986). Classificationssuch as been paid to thenomenclature
in whichsuch
those by Cronquist(1981) and Takhtajan(1980), ourknowledgeoftheearlyliterature
This situationhas in
althoughstillin frequentuse, have becomeoutdat- nameswereused is imperfect.
by Reveal's (1998)
thatbyGoldberg considerablepartbeen rectified
ed. Ofmorerecentclassifications,
is not
(1986) of the dicotyledonspredatesthe adventof Herculeanlabors. The principleof priority
fortaxa above the rank of family,almolecularstudiesat higherlevels, as does thatby mandatory
How- thoughauthorsare exhorted"generally"to follow
Dahlgrenet al. (1985) ofthemonocotyledons.
ever,the latterincorporatedmuch new data and thisprinciple(Greuteret al., 1994). We have tried
withgeneralusage whenassignschemesformanygroups. to balance priority
providedsynapomorphy
The recentsystemof Takhtajan(1997), although ingnamesto orders,buteven iffuturebibliographextremelyelaborate,is made less usefulbecause ic workdiscloses earlierordinalnames, changes
his propensityfor splittingoftenresultsin well- are notmandated.
Which clades should be recognizedin classifithenreassemknownfamiliesbeingdismembered,
thefindingsofrecent cation,or in our case, how should the ordersbe
bled as orders.Furthermore,
molecularstudies,despitebeingcited,have hardly circumscribed?Given the primaryprinciple of
that of recognizingclades and not
monophyly,
influencedhis classification.
We concludethatthereis a greatneed fora new, grades in classification,there are nevertheless
to be takenintoaccountwhen
plants,pro- manyconsiderations
offlowering
classification
phylogenetic
taxa at ordinalas well as all other
groups of circumscribing
viding names for major monophyletic
families.Obviously,it is notpossible,noris it de- hierarchicallevels above thatof species. Classifisirable,to name all clades in theentirephylogeny. cationis notonlya matterofgroupingaccordingto
but it is also a matter
wouldbe so cum- the principleof monophyly,
Anysuch completeclassification
of
philosophy
(notethatwhatever
bersomethatit wouldbe useless forgeneralcom- ofcommunication
munication.Systematistsneed to come to some namingis adopted,therehas to be someconsensus
whichclades to rec- as to the clades we are goingto use in general
kindofagreementconcerning
For us, this raises the
ognizeand name,so thata referencetoolof broad botanicalcommunication).
and used to discuss di- questionof ranking,thatis, afterhavingselected
utilitycan be formulated
to be named,theyhave to
offlowering
plant clades in the phylogeny
versity.An ordinalclassification
familiesis hereproposedforthatpurpose(pp. 538- be assignedan appropriateplace in the hierarchy,
542). It recognizesa selectednumberofmonophy- in ourcase, familyand order(e.g., Backlund& K.
letic suprafamilialgroups,that is, clades in the Bremer,1998; Stevens,1998). In choosingbetween
it is desirableto recofflowering
plantsthatare supportedby alternativecircumscriptions
phylogeny
at least one, and oftenseveral,lines of evidence. ognize groupsthat are well supported.It is also
These are clades to whichwe findit usefulto refer usefulto selectgroupsthathave somekindofeasalabout higher- ily observed morphologicalsynapomorphies,
when we communicateinformation
at theordinallevel and
ofthe flowering
plants.
thoughthismaybe difficult
level interrelationships
Volume 85, Number4

1998
AngiospermPhylogenyGroup
OrdinalClassification
533
even sometimesat the familylevel. Synapomor- uous because thiswill alwaysbe an arbitrary
dephies also ofteninclude (sometimesexclusively) cision.
Takhtajan(1997) optedin favorof"smaller,more
anatomical,biochemical,and developmental
charnaturalfamiliesand orders,whichare morecoheracters.
wherecharactersare easily
Manyof our ordinalnames are alreadywell es- ent and better-defined,
tablishedand used in earlier classificationsand grasped,and whichare moresuitableforinformasystematictreatments.So far as they represent tion retrievaland phylogeneticstudies,including
monophyletic
groups,we retainwell-known
orders cladistic analyses (e.g., because it reduces polyin theinterest
ofpreserving
stability.
In othercases, morphiccodings)."However,thesize ofa grouphas
the size of the orderscomes into consideration. nothingto do withits "naturalness."For a smaller
However,what is reasonablybroad circumscrip- group,one will oftenbe able to say moreaboutall
members,and so the characters
tion? From the point of view of memorization
of of its constituent
names,groupsof2-6 or a fewmorewouldseem to may be moreeasily grasped.However,segregates
familieslike Rube ideal, and thereis evidencethatsystematists
in ofwell establishedmonophyletic
biaceae
or
Asteraceae
(Gentianales)
(Asterales)
the past have commonly
recognizedgroupsof this
would
be more
also
by
Takhtajan's
generalization
size (Stevens,1997). However,withthediscoveries
thesmallerthegroup,the
of new species, genera,and families,the sizes of natural;bythiscriterion,
genera,families,and ordershave increased,and morenaturalit will necessarilybe, so thereis no
to be derivedfrom"naturalness."
manyordersnowcomprise10-20 families,or even rankingcriterion
If
by
"more
natural"
is meant"has moresynapomore.Otherorderscontaina fewfamiliesonly,and
is incorrect;the number
morphies"
then
this,
too,
if thereare onlytwoor threefamiliesin an order,
is not connectedto the size of
"one is notfarfromleavingthefamiliesunplaced" of synapomorphies
val- the groupor the hierarchicallevel at whichit is
(Copeland,1957). Concernsaboutthedoubtful
recognized.
ue of recognizing
similarlysmall groupshave also
In our classification,
these considerations
have
been expressedby others(e.g., Burtt,1977). Nevhad littleimpact.The principleof monophyly
in
have
number
ertheless,we
chosento recognizea
of
combination
withthedesirability
ofmaintaining
alsmall ordersbecause these representclades for
ready well establishedand familiarentitieshas
whichmonophyly
and relationshipsare well suplargelyformedthe ordinalclassification.
Monofamported,and this betterconveysthe interrelationilial orders(and monogeneric
families)are avoided
ships of the familiesincludedratherthanleaving
as much as possible, minimizingredundancyin
themunclassifiedto order.
In a fewcases we have,however,
classification.
recIn general,we adopta broad circumscription
of
orders(Ceratophyllales,
ognizedsomemonofamilial
the orders.We recognize462 familiesand 40 orAcorales,Arecales)because theseare sistergroups
ders of flowering
plants. Cronquist(1981) recog- of morethanone otherorder.Hence, the families
nized 321 familiesand 64 orders,Thorne(1992) ofthesemonofamilial
orderscannotbe includedin
440 familiesand 69 orders,and Takhtajan(1997) any otherorderwithoutviolatingmonophyly.
no less than589 familiesin 232 orders.Ourwider
The principleofmonophyly
in combination
with
ordinalcircumscription
is notbecause finerdetails themandatory
usage ofthefamilycategory
(Greuter
of the phylogeny
withinthe ordersare as yet un- et al., 1994) may lead to the recognition
of many
will smallfamilies.For example,in Dipsacales, ifDipclear, but because we thinkthe classification
be moreusefulwitha limitednumberoflargeror- sacaceae and Valerianaceaeare to be retainedas
ders.As we developmorefirmly
supportedphylog- familiesseparatefromCaprifoliaceae,
theprinciple
enies withinand amongorders,groupsat the in- ofmonophyly
also ofDierrequirestherecognition
fraordinal and supraordinal levels can be villaceae,Linnaeaceae,and Morinaceae(Backlund
recognized.Hence we anticipatethattherewill be & K. Bremer,1998; Backlund& Pyck,1998). This
oftheor- is because each oftheselatterfamiliesis thesister
littleneed to changethe circumscription
ders recognizedhere, except forinclusionof yet groupof morethan one familyso theycannotbe
unassignedfamiliesofunknownsystematic
position mergedwith any other familywithoutviolating
of misplacedfamilies.Additional monophyly.
and the transfer
Similarconsiderations
applyat the orordersmayhave to be recognizedas the phyloge- dinal level. Unfortunately,
no absolute guidelines
netic relationshipsof families that are not yet as to reasonablepracticecan be offered,but we
placed are clarified.Discussion as to whethera simplyobservethatcautionis alwaysin order.
In othercases thereare small familiesthatmay
widelyaccepted monophyletic
groupshould be a
orfamilyis largelyvac- be reducedto synonymy
oftheirsistergroupifthe
superorder,
order,suborder,
534
Annals of the
latterconsistsofa singlefamily.Examplesare Ca- thatconceivedbyBremeret al. (1995, 1996, 1997)

bombaceae,whichmaybe mergedwithNymphae- and since 1996 available on the Internet(Bremer
is based on various
aceae, and Kingdoniaceae,whichmay be merged et al., 1998). This classification
withCircaeasteraceae(Ranunculales).Such com- recentlypublishedmostlymolecularphylogenetic
maybe analyses (e.g., Chase et al., 1993; Chase et al.,
monlyrecognizedfamiliesthatnevertheless
mergedwiththeirsisterfamilyare in our classifi- 1995; Bremeret al., 1994; Struweet al., 1994; Nacation placed withinsquare bracketsbelow the dot et al., 1995; Nickrent& Soltis,1995; Soltiset
familywithwhichtheymay be merged(in Ran- al., 1995; Gadek et al., 1996; Gustafssonet al.,
unculaleseitherFumariaceaeor bothFumariaceae 1996; Mortonet al., 1996; Soltis & Soltis, 1997;
maybe mergedwithPapav- Soltis et al., 1997; Anderberget al., 1998; Backand Pteridophyllaceae
or lund & B. Bremer,1998; Bakkeret al., 1998; KaleitherPteridophyllaceae
eraceae; alternatively,
may be lersj6et al., 1998; Soltiset al., 1998; Thulinet al.,
both Fumariaceae and Pteridophyllaceae
referencesabove). The majordiffer1998; further
retainedas distinct).
revise family ences are in the expansionof Alismatales(includWe do not attemptto thoroughly
(includingDroIn generalwe followrecentau- ing also Araceae), Caryophyllales
circumscriptions.
thorsand attemptto recognizeas manymonophy- seraceae, Nepenthaceae,Polygonaceae,Plumbagileticfamiliesas possible.It shouldbe emphasized, naceae, and severalotherfamiliesoutsidethe traof a
the recognition
however,that followingadditional investigation ditional,core Caryophyllales),
widely circumscribedRosales (insomefamilieslistedbelowmaybe showntobe non- comparatively
eitherby cluding Rhamnaceae,Urticaceae,Moraceae, and
monophyletic;
revised circumscriptions,
intomonophyletic
taxaare not theirallies), in theadditionofa numberofsmaller
merging
or splitting,
Acorales,Arecales,Proteyet possible given our currentknowledge.Exam- orders(Ceratophyllales,
ples are Euphorbiaceaeand FlacourtiaceaeofMal- ales, Garryales,Aquifoliales),and in the deletion
BroNymphaeales,
pighiales(Kdllersj6et al., 1998) and severalfami- ofa fewothers(Aristolochiales,
Zygophyllales).Monolies of Myrtales(Contiet al., 1996; Gadek et al., meliales, Trochodendrales,
1996) and core Caryophyllales(which comprise cots and eudicots are not formallyranked and
Bas- named because it is not yet clear at whichlevel
Achatocarpaceae,Aizoaceae,Amaranthaceae,
ellaceae, Cactaceae, Caryophyllaceae,Didierea- theyshouldbe recognized.The same problemsocderived
ceae, Molluginaceae,Nyctaginaceae,Phytolacca- cur withcommelinoids(a phylogenetically
ceae,
Portulacaceae, Sarcobataceae, and subgroupofmonocots)and withrosidsand asterids
Stegnospermataceae; Hershkovitz & Zimmer, (subgroupsof eudicots),althoughthese are comfamilies monlyknownas subclasses Commelinidae,Rosi1997). Otherprobablynon-monophyletic
are Boragina- dae, and Asteridae,respectively.
thatcannotyet be recircumscribed
are
ceae (euasteridsI; Chase et al., 1993), Scrophular- Well supportedordinal interrelationships
amongthe
iaceae (Lamiales; Olmstead& Reeves, 1995), and shownin Figure 1. Interrelationships
Santalaceae (Santalales; Nickrent& Duff,1996; basal branchesof the treeand the positionof the
remaineluplant phylogeny
Nickrentet al., 1998). Brassicaceae (Brassicales) rootof the flowering
include also the former,
paraphyletic
Capparaceae sive. Withinthe eudicotsthereis increasingsuppen(Brassicaceae sensu strictobeing nested inside portfora large subgroupwithpredominantly
the core eudicots,
Capparaceae; Judd et al., 1994; Rodman et al., tamerousand isomerousflowers,
Santalales,SaxCaryophyllales,
1996). A supposedlyparallel case comprisesApi- mainlycomprising
aceae and Araliaceae (Apiales), since the former ifragales,rosids,and asterids.Rosids and asterids
have been assumed to be nestedinside the latter each comprisetwolargesubgroups,eurosidsI and
of II and euasteridsI and II, also receivingincreasing
(Plunkettet al., 1996). However,witha transfer
These correspondto the
fromApiaceae to Araliaceae, it supportas monophyletic.
Hydrocotyloideae
familiescan be rec- similarlynumberedrosid and asterid clades of
seems thattwomonophyletic
ognized, only a few genera remainingunplaced Chase et al. (1993).
Undereach ofthe supraordinalgroupsofmono(Plunkettet al., 1997). Delimitationof Bombacaceae, Malvaceae, Sterculiaceae, and Tiliaceae cots, commelinoids,core eudicots, rosids, etc.,
(Malvales) is problematical,and only Malvaceae thereare a numberof familieslisted withoutasto order.These familiesare knownto beare monophyletic
(Alversonet al., 1998; Bayeret signment
al., 1999). Here all fourare treatedtogetheras a long withinthe majorgroupunderwhichtheyare
family,Malvaceae sensu lato listed,but theirordinalpositionis stilluncertain.
single monophyletic
Similarly,Amborellaceae,Austrobaileyaceae,Ca1997).
(Judd& Manchester,
of nellaceae, etc.,are listedat thebeginningbecause
Our proposedclassificationis a modification
Volume85, Number4
1998
535
Angiosperm
Phylogeny
Group
100/75 Ceratophyllales
Laurales
10/S
5/-
05/BB
100/100
PW
(a
Cn
Piperales
sue
|
Magnoliales
Acorales
/855
Alismatales
56/50
~~~Asparagales
9 /100
8 ADioscoreales
Liliales
84/97
Pandanales
00/08
~~~~~~~Arecales
r-+.
05/-
oPoales
commelinoids
100/98
Zingiberales
CD
0|
T Ranunculales
28/88
85/Proteales
Cn _
100/99
Caryophyllales
San100/9
BB/-Santalales
CD
8:
Saxifragales
.
|95BGeraniales
SMalpighiales
o-,
Cn
COxalidales
77955810/-
00/58
0
CD.
12!!
:-
CD
CD
C
0
Cn
/ster/
F8/b5
f" Eriales
6~~~~0/-
Rosales
]
Cucurbitales
SolFagales
1
quiiMyrtales
100/94Brassicales
Ieu sdsI
euaseids 11
Males
~~~~~~~~Sapindales
98/~~~~~Cornales
Er98Dipsacales
Figure
1.0/Phyloge50/09
f
100/99Garryales
BobGn
naes
euasterids I
citedin thetext.Jackknife
support
is givenon thebranches
(a dashforvalues< 50%),first
jackknife
valuesfrom
analysisof 545 sequences oftherbcL,atpB,and 18S rDNA genes (D. E. Soltis,M. W. Chase, P. S. Soltis,D. Albach,
M. E. Mort,V. Savolainen,M. Zanis & J. S. Earris,unpublished,in prep.)and secondjackknifevalues fromanalysis
of 2538 rbcLsequences (Kallersjoet al., 1998).
536
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Wimpee,J. F. Smith,G. R. Furnier,S. H. Strauss,Q.Y. Xiang,G. M. Plunkett,P. S. Soltis,S. M. Swensen,
S. E. Williams,P. A. Gadek,C. J.Quinn,L. E. Eguiarte,
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Introduction
to Phylogeny
and Systematics
ofFlowering Greuter,W, F. R. Barrie,H. M. Burdet,W. G. Chaloner,
V. Demoulin,D. L. Hawksworth,
P. M. J0rgensen,
D. H.
Plants. 1st,2nd, 3rd eds. Compendium,
Uppsala UniNicolson,P. C. Silva, P. Trehane& J. McNeill. 1994.
versity,
Uppsala.
International
Code ofBotanicalNomenclature.
. 1998. Classification
Regnum
&
offlowVeg. 131.
eringplants. Internethttp://www.systbot.uu.se/classifiGustafsson,M. H. G., A. Backlund& B. Bremer.1996.
cation/overview.html.
Phylogenyof the Asteralessensu lato based on rbcL
Burtt,B. L. 1977. Classification
above the genus,as exsequences with particularreferenceto the Goodeniemplifiedby Gesneriaceae,withparallels fromother
aceae. P1. Syst.Evol. 199: 217-242.
groups.In K. Kubitzki(editor),Floweringplants:Evolutionand classification
M. A. & E. A. Zimmer.1997. On the evoof highercategories.PI. Syst. Hershkovitz,
Evol. Suppl. 1: 97-109.
lutionary
originsofthe cacti. Taxon46: 217-232.
Cantino,P. D., R. G. Olmstead& S. J. Wagstaff.
1997. A Heywood,V. H. 1977. Principlesand conceptsin theclascomparisonofphylogenetic
nomenclature
withthecursification
of highertaxa. In K. Kubitzki(editor),Flowrentsystem:A botanicalcase study.Syst.Biol. 46: 313eringplants:Evolutionand classification
ofhighercat331.
egories.P1. Syst.Evol. Suppl. 1: 1-12.
Chase, M. W., D. E. Soltis,R. G. Olmstead,D. Morgan, Judd,W S. & S. R. Manchester.1997. Circumscription
theybelongneitherin any of the phylogenetically

"basal" ordersat thebeginning
norin themonocots
or eudicots. Furthermore,
familieslisted directly
undermonocotswithoutan orderare monocotsbut
notcommelinoids,
and familiessimilarlylisteddirectlyundereudicotsand core eudicotsare eudicots or core eudicots,respectively,
butneitherrosids nor asterids.At the end of the systemis an
additionallist of familiesof uncertainposition.
Mostoftheseare probablyeudicots(includingcore
eudicots,rosids,and asterids),but so farthereare
no firmdata supporting
theirplacementanywhere
withinthe eudicots.
Volume 85, Number4

1998
bya preliminary
ofMalvaceae(Malvales)as determined
anatomical,palycladistic analysis of morphological,
49: 384nological,and chemicalcharacters.Brittonia
405.
, R. W. Sanders& M. J. Donoghue.1994. Angiophylogenetic
analyses.
spermfamilypairs: Preliminary
HarvardPap. Bot. 5: 1-51.
Kallersj6,M., J. S. Farris,M. W. Chase, B. Bremer,M. F.
Fay, C. J. Humphries,G. Petersen,0. Seberg & K.
jackknifeanalBremer.1998. Simultaneousparsimony
ysis of 2538 rbcL DNA sequences revealssupportfor
majorclades of greenplants,land plants,seed plants,
and flowering
plants.PI. Syst.Evol. [in press].
Liden, M., B. Oxelman,A. Backlund,L. Andersson,B.
Bremer,R. Eriksson,R. Moberg,I. Nordal,K. Persson,
M. Thulin& B. Zimmer.1997. Charlieis our darling.
Taxon46: 735-738.
H. 1977. Summarylecture.In K. Kubitzki
MerxmUller,
(editor),Floweringplants:Evolutionand classification
ofhighercategories.PI. Syst.Evol. Suppl. 1: 397-405.
Morton,
C. M., M. W.Chase, K. A. Kron& S. M. Swensen.
of the
1996. A molecularevaluationof the monophyly
orderEbenales based upon rbcL sequence data. Syst.
Bot. 21: 567-586.
Nadot,S., G. Bittar,L. Carter,R. Lacroix& B. Lejeune.
based
analysisofmonocotyledons
1995. A phylogenetic
and a new
on thechloroplast
generps4usingparsimony
numericalpheneticsmethod.Molec. Phylog.Evol. 4:
257-282.
Nickrent,D. L. & J. R. Duff.1996. Molecularstudiesof
parasiticplantsusingribosomalRNA. Pp. 28-52 in M.
T Moreno,J. I. Cubero,D. Berner,D. Joel,L. J. Musselman & C. Parker(editors),Advances in Parasitic
PlantResearch.Juntade Andalucfa,Direcci6nGeneral
de Investigaci6n
Agraria,C6rdoba,Spain.
ofangiosperm
& D. E. Soltis.1995. A comparison
fromnuclear18S rDNAand rbcLsequencphylogenies
es. Ann. MissouriBot. Gard.82: 208-234.
, J. R. Duff,A. E. Colwell,A. D. Wolfe,N. D.
Young,K. E. Steiner& C. W. dePamphilis.1998. Moand evolutionary
studiesof paralecular phylogenetic
siticplants.Pp. 211-241 in D. E. Soltis,P. S. Soltis&
ofPlantsII:
J.J. Doyle (editors),MolecularSystematics
DNA Sequencing.Kluwer,Boston.
Olmstead,R. G. & P. A. Reeves. 1995. Evidenceforthe
oftheScrophulariaceaebased on chloroplast
polyphyly
rbcLand ndhFsequences.Ann.MissouriBot.Gard.82:
176-193.
Plunkett,G. M., D. E. Soltis& P. S. Soltis.1996. Higher
level relationships
ofApiales(Apiaceaeand Araliaceae)
based on phylogeneticanalysis of rbcL sequences.
Amer.J. Bot. 83: 399-415.
. 1997. Classificationof the
&
betweenApiaceae and Araliaceaebased on
relationship
matK and rbcLsequence data. Amer.J. Bot. 84: 565580.

Reveal, J. L. 1998. Indices nominumsupragenericorum
plantarumvascularium.Internethttp://www.infoim.umd.
edu/PBIO/.WWW/supragen.html.
Rodman,J. E., K. G. Karol,R. A. Price & K. J. Sytsma.
and Dahlgren'sexpanded
1996. Molecules,morphology,
orderCapparales.Syst.Bot. 21: 289-307.
relationSoltis,D. E. & P. S. Soltis. 1997. Phylogenetic
ships in Saxifragaceaesensu lato: A comparisonoftopologies based on 18S rDNA and rbcL sequences.
Amer.J. Bot. 84: 504-522.
D. R. Morgan,S. M. Swensen,B. C.
Mullin,J. M. Dowd & P. G. Martin.1995. Chloroplast
gene sequence data suggesta singleoriginof the predispositionfor symbioticnitrogenfixationin angiosperms.Proc. Natl.Acad. Sci., U.S.A. 92: 2647-2651.
, M. Mort,M. W. Chase, V. Savolainen,
S. B. Hoot & C. M. Morton.1998. Inferring
complex
phylogeniesusing parsimony:An empiricalapproach
Syst.
usingthreelargeDNA data sets forangiosperms.
Biol. 47: 32-42.
, D. L. Nickrent,L. A. Johnson,W. J.
9
Hahn, S. B. Hoot,J. A. Sweere,R. K. Kuzoff,K. A.
Kron,M. W. Chase, S. M. Swensen,E. A. Zimmer,C.
Shu-Miaw,L. J. Gilliespie,W. J. Kress& K. J. Sytsma.
inferredfrom18S ribo1997. Angiospermphylogeny
somal DNA sequences. Ann. MissouriBot. Gard. 84:
1-49.
in botany,
classification
Stevens,P. F. 1986. Evolutionary
1960-1985. J. ArnoldArbor.67: 313-339.
botanicalclassifications:
. 1997. How to interpret
Bioscience47: 250-250.
Suggestionsfromhistory.
. 1998. What kind of classificationshould the
use to be saved? Pp. 295-319 in
practisingtaxonomist
M. J.E. Coode & D. A. Simpson(editors),
J.Dransfield,
Plant Diversityin Malesia III. Royal BotanicGardens,
Kew.
Struwe,L., V. A. Albert& B. Bremer.1994. Cladistics
of the Gentianales.Claand familylevel classification
distics10: 175-206.
offlowTakhtajan,A. 1980. Outlineof the classification
Bot. Rev. 46: 225-359.
eringplants(Magnoliophyta).
of Flowering
. 1997. Diversityand Classification
Plants.ColumbiaUniv.Press,New York.
of the
classification
Thorne,R. F. 1976. A phylogenetic
Evol. Biol. 9: 35-106.
Angiospermae.
classification
of
. 1992. An updatedphylogenetic
flowering
plants.Aliso 13: 365-389.
Thulin,M., B. Bremer,
J. Richardson,J. Niklasson,M. F.
ofthe
Fay & M. W. Chase. 1998. Familyrelationships
enigmaticrosidgeneraBarbeyaand Dirachmafromthe
Hornof Africaregion.P1. Syst.Evol. 213: 103-119.
537
Annals of the
538
CLASSIFICATION OF FLOWERING PLANTS
Amborellaceae
Austrobaileyaceae
Canellaceae
Chloranthaceae
Hydnoraceae
Illiciaceae
Nymphaeaceae
[+ Cabombaceae]
Rafflesiaceae
Schisandraceae
Trimeniaceae
Winteraceae
Bisch.
Ceratophyllales
Ceratophyllaceae
Laurales Perleb
Atherospermataceae
Calycanthaceae
Gomortegaceae
Hernandiaceae
Lauraceae
Monimiaceae
Siparunaceae
MagnolialesBromhead
Annonaceae
Degeneriaceae
Eupomatiaceae
Himantandraceae
Magnoliaceae
Myristicaceae
PiperalesDumort.
Aristolochiaceae
Lactoridaceae
Piperaceae
Saururaceae
Limnocharitaceae
Posidoniaceae
Potamogetonaceae
Ruppiaceae
Scheuchzeriaceae
Tofieldiaceae
Zosteraceae
AsparagalesBromhead
Agapanthaceae
Agavaceae
Alliaceae
Amaryllidaceae
Anemarrhenaceae
Anthericaceae
Aphyllanthaceae
Asparagaceae
Asphodelaceae
Asteliaceae
Behniaceae
Blandfordiaceae
Boryaceae
Convallariaceae
Doryanthaceae
Hemerocallidaceae
Herreriaceae
Hesperocallidaceae
Hyacinthaceae
Hypoxidaceae
Iridaceae
Ixioliriaceae
Lanariaceae
Laxmanniaceae
Orchidaceae
Tecophilaeaceae
Themidaceae
Xanthorrhoeaceae
Xeronemataceae
MONOCOTS
Corsiaceae
Japonoliriaceae
Nartheciaceae
Petrosaviaceae
Triuridaceae
DioscorealesHook. f.
Burmanniaceae
Dioscoreaceae
Taccaceae
Thismiaceae
Trichopodaceae
AcoralesReveal
Acoraceae
Liliales Perleb
Alstroemeriaceae
Campynemataceae
Colchicaceae
Liliaceae
Luzuriagaceae
Melanthiaceae
Philesiaceae
Ripogonaceae
Smilacaceae
AlismatalesDumort.
Alismataceae
Aponogetonaceae
Araceae
Butomaceae
Cymodoceaceae
Hydrocharitaceae
Juncaginaceae
Volume 85, Number4

1998
(cont'd.)
PandanalesLindl.
Cyclanthaceae
Pandanaceae
Stemonaceae
Velloziaceae
COMMELINOIDS
Abolbodaceae
Bromeliaceae
Dasypogonaceae
Hanguanaceae
Mayacaceae
Rapateaceae
ArecalesBromhead
Arecaceae
CommelinalesDumort.
Commelinaceae
Haemodoraceae
Philydraceae
Pontederiaceae
Poales Small
Anarthriaceae
Centrolepidaceae
Cyperaceae
Ecdeiocoleaceae
Eriocaulaceae
Flagellariaceae
Hydatellaceae
Joinvilleaceae
Juncaceae
Poaceae
Prioniaceae
Restionaceae
Sparganiaceae
Thurniaceae
Typhaceae
Xyridaceae
ZingiberalesGriseb.
Cannaceae
Costaceae
Heliconiaceae
Lowiaceae
Marantaceae
Musaceae
Strelitziaceae
Zingiberaceae
EUDICOTS
Buxaceae
Didymelaceae
Sabiaceae
Trochodendraceae
[+Tetracentraceae]
ProtealesDumort.
Nelumbonaceae
Platanaceae
Proteaceae
RanunculalesDumort.
Berberidaceae
Circaeasteraceae
[+Kingdoniaceae]
Eupteleaceae
Lardizabalaceae
Menispermaceae
Papaveraceae
[+ Fumariaceae]
[+Pteridophyllaceae]
Ranunculaceae
CORE EUDICOTS
Aextoxicaceae
Berberidopsidaceae
Dilleniaceae
Gunneraceae
Myrothamnaceae
Vitaceae
Perleb
Caryophyllales
Achatocarpaceae
Aizoaceae
Amaranthaceae
Ancistrocladaceae
Asteropeiaceae
Basellaceae
Cactaceae
Caryophyllaceae
Didiereaceae
Dioncophyllaceae
Droseraceae
Drosophyllaceae
Frankeniaceae
Molluginaceae
Nepenthaceae
Nyctaginaceae
Physenaceae
Phytolaccaceae
Plumbaginaceae
Polygonaceae
Portulacaceae
Rhabdodendraceae
Sarcobataceae
Simmondsiaceae
Stegnospermataceae
Tamaricaceae
SantalalesDumort.
Olacaceae
Opiliaceae
539
540
Annals of the
(cont'd.)
Loranthaceae
Misodendraceae
Santalaceae
SaxifragalesDumort.
Altingiaceae
Cercidiphyllaceae
Crassulaceae
Daphniphyllaceae
Grossulariaceae
Haloragaceae
Hamamelidaceae
Iteaceae
Paeoniaceae
Penthoraceae
Pterostemonaceae
Saxifragaceae
Tetracarpaeaceae
ROSIDS
Aphloiaceae
Crossosomataceae
Ixerbaceae
Krameriaceae
Picramniaceae
Podostemaceae
Stachyuraceae
Staphyleaceae
Tristichaceae
Zygophyllaceae
GeranialesDumort.
Francoaceae
Geraniaceae
[+ Hypseocharitaceae]
Greyiaceae
Ledocarpaceae
Melianthaceae
Vivianiaceae
EUROSIDS I
Celastraceae
Huaceae
Parnassiaceae
[ + Lepuropetalaceae]
Stackhousiaceae
CucurbitalesDumort.
Anisophylleaceae
Begoniaceae
Coriariaceae
Corynocarpaceae
Cucurbitaceae
Datiscaceae
Tetramelaceae
Fabales Bromhead
Fabaceae
Polygalaceae
Quillajaceae
Surianaceae
Fagales Engl.
Betulaceae
Casuarinaceae
Fagaceae
Juglandaceae
Myricaceae
Nothofagaceae
Rhoipteleaceae
Ticodendraceae
MalpighialesMart.
Achariaceae
Balanopaceae
Caryocaraceae
Chrysobalanaceae
Clusiaceae
Dichapetalaceae
Erythroxylaceae
Euphorbiaceae
Euphroniaceae
Flacourtiaceae
Goupiaceae
Hugoniaceae
Humiriaceae
Irvingiaceae
Ixonanthaceae
Lacistemataceae
Linaceae
Malesherbiaceae
Malpighiaceae
Medusagynaceae
Ochnaceae
Pandaceae
Passifloraceae
Putranjivaceae
Quiinaceae
Rhizophoraceae
Salicaceae
Scyphostegiaceae
Trigoniaceae
Turneraceae
Violaceae
Oxalidales Heintze
Cephalotaceae
Connaraceae
Cunoniaceae
Elaeocarpaceae
Oxalidaceae
Tremandraceae
Volume 85, Number4

1998
(cont'd.)
Rosales Perleb
Barbeyaceae
Cannabaceae
Cecropiaceae
Celtidaceae
Dirachmaceae
Elaeagnaceae
Moraceae
Rhamnaceae
Rosaceae
Ulmaceae
Urticaceae
EUROSIDS II
Tapisciaceae
BrassicalesBromhead
Akaniaceae
[ + Bretschneideraceae]
Bataceae
Brassicaceae
Caricaceae
Emblingiaceae
Gyrostemonaceae
Koeberliniaceae
Limnanthaceae
Moringaceae
Pentadiplandraceae
Resedaceae
Salvadoraceae
Setchellanthaceae
Tovariaceae
Tropaeolaceae
MalvalesDumort.
Bixaceae
[+Diegodendraceae]
Cistaceae
Cochlospermaceae
Dipterocarpaceae
Malvaceae
Muntingiaceae
Neuradaceae
Sarcolaenaceae
Sphaerosepalaceae
Thymelaeaceae
MyrtalesRchb.
Alzateaceae
Combretaceae
Crypteroniaceae
Heteropyxidaceae
Lythraceae
Melastomataceae
Memecylaceae
Myrtaceae
Oliniaceae
Onagraceae
Penaeaceae
Psiloxylaceae
Rhynchocalycaceae
Vochysiaceae
Sapindales Dumort.
Anacardiaceae
Biebersteiniaceae
Burseraceae
Kirkiaceae
Meliaceae
Nitrariaceae
[+ Peganaceae]
Rutaceae
Sapindaceae
Simaroubaceae
ASTERIDS
CornalesDumort.
Cornaceae
[+ Nyssaceae]
Grubbiaceae
Hydrangeaceae
Hydrostachyaceae
Loasaceae
Ericales Dumort.
Actinidiaceae
Balsaminaceae
Clethraceae
Cyrillaceae
Diapensiaceae
Ebenaceae
Ericaceae
Fouquieriaceae
Halesiaceae
Lecythidaceae
Marcgraviaceae
Myrsinaceae
Pellicieraceae
Polemoniaceae
Primulaceae
Roridulaceae
Sapotaceae
Sarraceniaceae
Styracaceae
Symplocaceae
Ternstroemiaceae
Tetrameristaceae
Theaceae
Theophrastaceae
EUASTERIDS I
Boraginaceae
Plocospermataceae
Vahliaceae
541
Annals of the
542
(cont'd.)
GarryalesLindl.
Aucubaceae
Eucommiaceae
Garryaceae
Oncothecaceae
GentianalesLindl.
Apocynaceae
Gelsemiaceae
Gentianaceae
Loganiaceae
Rubiaceae
Lamiales Bromhead
Acanthaceae
Avicenniaceae
Bignoniaceae
Buddlejaceae
Byblidaceae
Cyclocheilaceae
Gesneriaceae
Lamiaceae
Lentibulariaceae
Myoporaceae
Oleaceae
Orobanchaceae
Paulowniaceae
Pedaliaceae
[+ Martyniaceae]
Phrymaceae
Plantaginaceae
Schlegeliaceae
Scrophulariaceae
Stilbaceae
Tetrachondraceae
Verbenaceae
Solanales Dumort.
Convolvulaceae
Hydroleaceae
Montiniaceae
Solanaceae
Sphenocleaceae
EUASTERIDS II
Adoxaceae
Bruniaceae
Carlemanniaceae
Columelliaceae
[+Desfontainiaceae]
Eremosynaceae
Escalloniaceae
Icacinaceae
Polyosmaceae
Sphenostemonaceae
Tribelaceae
Apiales Nakai
Apiaceae
Araliaceae
Aralidiaceae
Griseliniaceae
Melanophyllaceae
Pittosporaceae
Torricelliaceae
AquifolialesSenft
Aquifoliaceae
Helwingiaceae
Phyllonomaceae
AsteralesLindl.
Alseuosmiaceae
Argophyllaceae
Asteraceae
Calyceraceae
Campanulaceae
[+Lobeliaceae]
Carpodetaceae
Donatiaceae
Goodeniaceae
Menyanthaceae
Pentaphragmataceae
Phellinaceae
Rousseaceae
Stylidiaceae
Dipsacales Dumort.
Caprifoliaceae
Diervillaceae
Dipsacaceae
Linnaeaceae
Morinaceae
Valerianaceae
FAMILIES OF UNCERTAIN POSITION
Balanophoraceae
Bonnetiaceae
Cardiopteridaceae
Ctenolophonaceae
Cynomoriaceae
Cytinaceae
Dipentodontaceae
Elatinaceae
Geissolomataceae
Hoplestigmataceae
Kaliphoraceae
Lepidobotryaceae
Lissocarpaceae
Lophopyxidaceae
Medusandraceae
Metteniusaceae
Mitrastemonaceae
Paracryphiaceae
Volume 85, Number4

1998
(cont'd.)
Pentaphylacaceae
Peridiscaceae
Plagiopteraceae
Pottingeriaceae
Sladeniaceae
Strasburgeriaceae
Tepuianthaceae
ORDINAL SYNONYMS
AcanthalesLindl.
= Lamiales
Acerales Lindl.
= Sapindales
ActinidialesTakht.ex Reveal
= Ericales
AdoxalesNakai
- notaccepted,familyunder
euasteridsII
Aesculales Bromhead
Asarales Horan.
= Piperales
AsclepiadalesDumort.
= Gentianales
AstelialesDumort.
= Asparagales
AtriplicalesHoran.
= Caryophyllales
AucubalesTakht.
= Garryales
Austrobaileyales
Takht.ex Reveal
- notaccepted,familyat beginning
of system
AvenalesBromhead
= Poales
BalanopalesEngl.
= Malpighiales
= Sapindales
BalanophoralesDumort.
- notaccepted,familyunplaced
BalsaminalesLindl.
= Asparagales
BarbeyalesTakht.& Reveal
AgavalesHutch.
Alliales Traub
= Asparagales
Hutch.
Alstroemeriales
= Liliales
AltingialesDoweld
= Saxifragales
= Ericales
= Rosales
Batales Engl.
= Brassicales
BegonialesDumort.
= Cucurbitales
BerberidalesDumort.
= Ranunculales
Dumort.
Amaranthales
BetulalesBromhead
Bromhead
Amaryllidales
Biebersteiniales
Takht.
AmbrosialesDumort.
BignonialesLindl.
AmmialesSmall
Bixales Lindl.
AmomalesLindl.
Arales Dumort.
BoraginalesDumort.
euasteridsI
BrexialesLindl.
eurosidsI
BromelialesDumort.
commelinoids
BrunialesDumort.
euasteridsII
BrunonialesLindl.
AralialesReveal
BurmannialesHeintze
AralidialesTakht.ex Reveal
BurseralesBaskerville
Aristolochiales
Dumort.
ButomalesHutch.
= Caryophyllales
= Asparagales
= Asterales
= Apiales
= Zingiberales
Ancistrocladales
Takht.
= Caryophyllales
AnnonalesLindl.
= Magnoliales
AnthobolalesDumort.
= Santalales
ApocynalesBromhead
= Gentianales
Hutch.
Aponogetonales
= Alismatales
= Alismatales
= Apiales
= Apiales
= Piperales
= Fagales
= Sapindales
= Lamiales
= Malvales
= Asterales
= Dioscoreales
= Sapindales
= Alismatales
543
544
Annals of the
(cont'd.)
Buxales Takht.ex Reveal

eudicots
ByblidalesNakai ex Reveal
= Lamiales
Cactales Dumort.
= Caryophyllales
CallitrichalesDumort.
= Lamiales
CalycanthalesMart.
= Laurales
CalyceralesTakht.ex Reveal
= Asterales
CampanulalesRchb.
= Asterales
Canellales Cronquist
of system
Cannales Dumort.
= Zingiberales
CapparalesHutch.
= Brassicales
CaprifolialesLindl.
= Dipsacales
Takht.
Cardiopteridales
euasteridsII
CardualesSmall
= Asterales
Caricales L. D. Benson
= Brassicales
Cassiales Horan.
= Fabales
CasuarinalesLindl.
= Fagales
CelastralesBaskerville
eurosidsI
Centrolepidales
Takht.
= Poales
CephalotalesNakai
= Oxalidales
Hu ex Reveal
Cercidiphyllales
= Saxifragales
ChenopodialesDumort.
= Caryophyllales
ChironialesGriseb.
= Gentianales
A. C. Sm. ex J. -F. Leroy

Chloranthales
of system
CinchonalesLindl.
= Gentianales
CircaeasteralesTakht.
= Ranunculales
CistalesRchb.
= Malvales
CitralesDumort.
= Sapindales
Cocosales Nakai
= Arecales
ColchicalesDumort.
= Liliales
CombretalesBaskerville
= Myrtales
ConnaralesTakht.ex Reveal
= Cunoniales
ConvolvulalesDumort.
= Solanales
CoriarialesLindl.
= Cucurbitales
CorylalesDumort.
= Fagales
Takht.
Corynocarpales
= Cucurbitales
CrassulalesLindl.
= Saxifragales
Crossosomatales
Takht.ex Reveal
- notaccepted,familyunderrosids
CunonialesHutch.
= Oxalidales
CyclanthalesJ. H. Schaffn.
= Pandanales
CymodocealesNakai
= Alismatales
Burnett
Cynomoriales
CyperalesHutch.
= Poales
CytinalesDumort.
Daphnales Lindl.
= Malvales
DaphniphyllalesPulle ex Cronquist
= Saxifragales
Datiscales Dumort.
= Cucurbitales
Desfontainiales
Takht.
euasteridsII
Diapensiales Engl. & Gilg
= Ericales
DidymelalesTakht.
eudicots
Dilleniales Hutch.
- notaccepted,familyundercore
eudicots
DioncophyllalesTakht.ex Reveal
= Caryophyllales
DiospyralesPrantl
= Ericales
DroseralesGriseb.
= Caryophyllales
Volume 85, Number4
1998
(cont'd.)
Ebenales Engl.
= Ericales
Elaeagnales Bromhead
= Rosales
ElaeocarpalesTakht.
= Oxalidales
ElatinalesNakai
Elodeales Nakai
= Alismatales
EmpetralesRaf.
= Ericales
EriocaulalesNakai
= Poales
EucommialesNemejc ex Cronquist
= Garryales
EuphorbialesLindl.
= Malpighiales
EupomatialesTakht.ex Reveal
= Magnoliales
EuptelealesHu ex Reveal
= Ranunculales
EuryalalesH.L.Li
of system
Ficales Dumort.
= Rosales
FlacourtialesHeintze
= Malpighiales
FouquierialesTakht.ex Reveal
= Ericales
FrancoalesTakht.
= Geraniales
FrangulalesWirtg.
= Rosales
Galiales Bromhead
= Gentianales
GeissolomatalesTakht.ex Reveal
GesnerialesDumort.
= Lamiales
GlaucidialesTakht.ex Reveal
= Ranunculales
GlobularialesDumort.
= Lamiales
GoodenialesLindl.
= Asterales
GreyialesTakht.
= Geraniales
GrossularialesLindl.
= Saxifragales
GunneralesTakht.ex Reveal
eudicots
GyrocarpalesDumort.
= Laurales
Gyrostemonales
Takht.
= Brassicales
HaemodoralesHutch.
= Commelinales
HaloragalesBromhead
= Saxifragales
HamamelidalesGriseb.
= Saxifragales
HanguanalesR. Dahlgrenex Reveal
= notaccepted,familyunder
commelinoids
HelleboralesNakai
= Ranunculales
HelwingialesTakht.
= Aquifoliales
Himantandrales
Doweld & Shevyryova
= Magnoliales
HippuridalesPulle ex Reveal
= Lamiales
HomalialesBromhead
= Malpighiales
HortensialesGriseb.
= Cornales
HydatellalesCronquist
= Poales
HydnoralesTakht.ex Reveal
of system
HydrangealesNakai
= Cornales
Hydrastidales
Takht.
= Ranunculales
Hydropeltidales
(Bartl.)Spenn.
- notaccepted,family
Nymphaeaceaeat beginningof
system
Hydrostachyales
Diels ex Reveal
= Cornales
HypericalesDumort.
= Malpighiales
HypoxidalesTakht.
= Asparagales
Icacinales Tiegh.ex Reveal
euasteridsII
Illiciales Hu ex Cronquist
of system
IridalesRaf.
= Asparagales
Ixiales Lindl.
= Asparagales
JasminalesDumort.
= Lamiales
JuglandalesDumort.
= Fagales
545
546
Annals of the
(cont'd.)
JulianialesEngl.
= Sapindales
JuncaginalesHutch.
= Alismatales
JuncalesDumort.
= Poales
LacistematalesBaskerville
= Malpighiales
LactoridalesTakht.ex Reveal
= Piperales
LardizabalalesLoconte
= Ranunculales
LecythidalesCronquist
= Ericales
LeitnerialesEngl.
= Sapindales
LentibularialesLindl.
= Lamiales
LigustralesBarti.ex Bisch.
= Lamiales
LimnanthalesNakai
= Brassicales
Linales Baskerville
= Malpighiales
Loasales Bessey
= Cornales
LoganialesLindl.
= Gentianales
LoniceralesT. Liebe
= Dipsacales
LoranthalesDumort.
= Santalales
LythralesCaruel
= Myrtales
MarathralesDumort.
- not accepted, family
Podostemaceaeunderrosids
MayacalesNakai
commelinoids
MedusagynalesTakht.
= Malpighiales
MedusandralesBrenan
MelanthialesR. Dahlgrenex Reveal
= Liliales
MelastomatalesOliv.
= Myrtales
Meliales Lindl.
= Sapindales
MenispermalesBromhead
= Ranunculales
T. Yamaz. ex Takht.
Menyanthales
= Asterales
Metteniusales
Takht.
Mitrastemonales
Makino
MonimialesDumort.
= Laurales
MoringalesNakai
= Brassicales
MyricalesEngl.
= Fagales
Myristicales
Thome
= Magnoliales
Myrothamnales
Nakai ex Reveal
eudicots
MyrsinalesSpenn.
= Ericales
Najadales Dumort.
= Alismatales
NarcissalesDumort.
= Asparagales
NarthecialesReveal & Zomlefer
monocots
NelumbonalesReveal
= Proteales
NepenthalesDumort.
= Caryophyllales
NolanalesLindl.
= Solanales
NyctaginalesDumort.
= Caryophyllales
NymphaealesDumort.
= notaccepted,familyat beginning
of system
OchnalesHutch.ex Reveal
= Malpighiales
OenotheralesBromhead
= Myrtales
Olacales Benth.
= Santalales
Oleales Lindl.
= Lamiales
OnagralesRchb.
= Myrtales
OpuntialesWillk.
= Caryophyllales
OrchidalesRaf.
= Asparagales
Paeoniales Heintze
= Saxifragales
Pandales Engl. & Gilg
= Malpighiales
PapaveralesDumort.
= Ranunculales
Takht.
Paracryphiales
ParidalesDumort.
= Liliales
Volume 85, Number4

1998
(cont'd.)
ParnassialesNakai
eurosidsI
PassifloralesDumort.
= Malpighiales
Penaeales Lindl.
= Myrtales
PetiverialesLindl.
= Caryophyllales
PetrosavialesTakht.
monocots
PhilydralesDumort.
= Commelinales
PhysenalesTakht.
= Caryophyllales
PinguiculalesDumort.
= Lamiales
Pittosporales
Lindl.
= Apiales
PlantaginalesLindl.
= Lamiales
PlatanalesJ. H. Schaffn.
= Proteales
PlumbaginalesLindl.
= Caryophyllales
PodophyllalesDumort.
= Ranunculales
PodostemalesLindl.
= notaccepted,familyunderrosids
PolemonialesBromhead
= Ericales
PolygalalesDumort.
= Fabales
PolygonalesDumort.
= Caryophyllales
PontederialesHook. f.
= Commelinales
PortulacalesDumort.
= Caryophyllales
PosidonialesNakai
= Alismatales
Dumort.
Potamogetonales
= Alismatales
PrimulalesDumort.
= Ericales
QuercalesBurnett
= Fagales
RafflesialesOliv.
of system
Resedales Dumort.
= Brassicales
RestionalesJ. H. Schaffn.
= Poales
RhamnalesDumort.
= Rosales
RhinanthalesDumort.
= Lamiales
RhizophoralesTiegh.ex Reveal
= Malpighiales
RhodoralesHoran.
= Ericales
RhoiptelealesNovakex Reveal
= Fagales
RoridulalesNakai
= Ericales
Rubiales Dumort.
= Gentianales
RuppialesNakai
= Alismatales
RutalesPerleb
= Sapindales
Sabiales Takht.
= notaccepted,familyunder
eudicots
Salicales Lindl.
= Malpighiales
SalvadoralesR. Dahlgrenex Reveal
= Brassicales
SamolalesDumort.
= Ericales
SamydalesDumort.
= Malpighiales
SanguisorbalesDumort.
= Rosales
SapotalesHook. f.
= Ericales
SarracenialesBromhead
= Ericales
ScheuchzerialesB. Boivin
= Alismatales
ScleranthalesDumort.
= Caryophyllales
Lindl.
Scrophulariales
= Lamiales
Croizat
Scyphostegiales
= Malpighiales
Sedales Rchb.
= Saxifragales
Silenales Lindl.
= Caryophyllales
SimmondsialesReveal
= Caryophyllales
SmilacalesLindl.
= Liliales
StellarialesDumort.
= Caryophyllales
StylidialesTakht.ex Reveal
= Asterales
StyracalesBisch.
= Ericales
Taccales Dumort.
= Dioscoreales
547
Annals of the
548
(cont'd.)
TamalesDumort.
= Dioscoreales
TamaricalesHutch.
= Caryophyllales
TecophilaealesTraubex Reveal
= Asparagales
Theales Lindl.
= Ericales
TheligonalesNakai
= Gentianales
ThymelaealesWillk.
= Malvales
Tiliales Caruel
= Malvales
TofieldialesReveal & Zomlefer

= Alismatales
TorricellialesTakht.ex Reveal
= Apiales
TovarialesNakai
= Brassicales
TrillialesTakht.
= Liliales
TriuridalesHook. f.
- notaccepted,familyundermonocots
Trochodendrales
Takht.ex Cronquist
eudicots
TropaeolalesTakht.ex Reveal
= Brassicales
TurneralesDumort.
= Malpighiales
TyphalesDumort.
= Poales
UlmalesLindl.
= Rosales
UrticalesDumort.
= Rosales
VaccinialesDumort.
= Ericales
VallisnerialesNakai
= Alismatales
VellozialesR. Dahlgrenex Reveal

= Pandanales
VeratralesDumort.
= Liliales
VerbenalesHoran.
= Lamiales
ViburnalesDumort.
euasteridsII
Vincales Horan.
= Gentianales
Violales Perleb
= Malpighiales
VitalesReveal
eudicots
VochysialesDumort.
= Myrtales
WinteralesA. C. Sm. ex Reveal
of system
XyridalesLindl.
= Poales
ZosteralesNakai
= Alismatales
Takht.
Zygophyllales
- notaccepted,familyunderrosids
SELECTED FAMILIAL SYNONYMS
Abrophyllaceae
= Carpodetaceae
Acanthochlamydaceae
= Velloziaceae
Aceraceae
= Sapindaceae
Achradaceae
= Sapotaceae
Aegicerataceae
Myrsinaceae
Agdestidaceae
= Phytolaccaceae
Aitoniaceae
= Meliaceae
Alangiaceae
= Cornaceae
Aloaceae
= Asphodelaceae
Alsinaceae
= Caryophyllaceae
Ambrosiaceae
= Asteraceae
Amygdalaceae
= Rosaceae
Androstachyaceae
= Euphorbiaceae
Antoniaceae
= Loganiaceae
Apodanthaceae
= Rafflesiaceae
Apostasiaceae
= Orchidaceae
Aptandraceae
= Olacaceae
Aristoteliaceae
= Elaeocarpaceae
Asclepiadaceae
= Apocynaceae
Asteranthaceae
= Lecythidaceae
Averrhoaceae
= Oxalidaceae
Avetraceae
= Dioscoreaceae
Volume 85, Number4

1998
(cont'd.)
Balanitaceae
= Zygophyllaceae
Barbeiuaceae
= Phytolaccaceae
Barclayaceae
= Nymphaeaceae
Barringtoniaceae
= Lecythidaceae
Baueraceae
= Cunoniaceae
Baxteriaceae
= Dasypogonaceae
Bembiciaceae
= Flacourtiaceae
Berzeliaceae
= Bruniaceae
Bischofiaceae
= Euphorbiaceae
Blepharocaryaceae
= Anacardiaceae
Boerlagellaceae
= Sapotaceae
Bombacaceae
= Malvaceae
Boopidaceae
= Calyceraceae
Bretschneideraceae
= Akaniaceae
Brexiaceae
= Celastraceae
Brunelliaceae
= Cunoniaceae
Brunoniaceae
= Goodeniaceae
Bumeliaceae
= Sapotaceae
Burchardiaceae
= Colchicaceae
Byttneriaceae
= Malvaceae
Cabombaceae
= Nymphaeaceae
Caesalpiniaceae
= Fabaceae
Calectasiaceae
= Dasypogonaceae
Callitrichaceae
= Plantaginaceae
Calochortaceae
= Liliaceae
Camelliaceae
= Theaceae
Canotiaceae
= Celastraceae
Cansjeraceae
= Opiliaceae
Capparaceae
= Brassicaceae
Carduaceae
= Asteraceae
Cassythaceae
= Lauraceae
Chailletiaceae
= Dichapetalaceae
Chenopodiaceae
= Amaranthaceae
Chionographidaceae
= Melanthiaceae
Chloanthaceae
= Lamiaceae
Cichoriaceae
= Asteraceae
Cleomaceae
= Brassicaceae
Cneoraceae
= Rutaceae
Cobaeaceae
= Polemoniaceae
Compositae
= Asteraceae
Conostylidaceae
= Haemodoraceae
Cordiaceae
= Boraginaceae
Coridaceae
= Primulaceae
Corokiaceae
= Argophyllaceae
Corylaceae
= Betulaceae
Croomiaceae
= Stemonaceae
Cruciferae
= Brassicaceae
Curtisiaceae
= Comaceae
Cuscutaceae
= Convolvulaceae
Cyananthaceae
= Campanulaceae
Cyanastraceae
= Tecophilaeaceae
nom.illeg.
Cynocrambaceae
= Rubiaceae
Cyphiaceae
= Campanulaceae
Cyphocarpaceae
= Campanulaceae
Cypripediaceae
= Orchidaceae
Dactylanthaceae
= Balanophoraceae
549
550
Annals of the
(cont'd.)
Davidiaceae
Frangulaceae
Davidsoniaceae
Fumariaceae
Decaisneaceae
Funkiaceae
Desfontainiaceae
Galacaceae
Dialypetalanthaceae
Geitonoplesiaceae
Dianellaceae
Geniostomaceae
Dichondraceae
Geosiridaceae
Diclidantheraceae
Gisekiaceae
Diegodendraceae
Glaucidiaceae
Dionaeaceae
Globulariaceae
Dracaenaceae
Goetzeaceae
Duabangaceae
Gonystylaceae
= Comaceae
= Cunoniaceae
= Lardizabalaceae
= Columelliaceae
= Rubiaceae
= Hemerocallidaceae
= Convolvulaceae
= Polygalaceae
= Bixaceae
= Droseraceae
= Convallariaceae
= Lythraceae
Duckeodendraceae
= Solanaceae
Dulongiaceaenom.illeg.
= Phyllonomaceae
Dysphaniaceae
= Amaranthaceae
Ehretiaceae
= Boraginaceae
= Rhamnaceae
= Papaveraceae
= Agavaceae
= Diapensiaceae
= Hemerocallidaceae
= Loganiaceae
= Iridaceae
= Phytolaccaceae
= Ranunculaceae
= Plantaginaceae
= Solanaceae
= Thymelaeaceae
Gouaniaceae
= Rhamnaceae
Gramineae
= Poaceae
Gronoviaceae
= Loasaceae
Gustaviaceae
= Lecythidaceae
Ellisiophyllaceae
Guttiferae
Empetraceae
Gyrocarpaceae
Epacridaceae
Halophilaceae
Eremolepidaceae
Halophytaceae
= Scrophulariaceae
= Ericaceae
= Ericaceae
= Santalaceae
Eriospermaceae
= Convallariaceae
= Clusiaceae
= Hemandiaceae
= Hydrocharitaceae
= Amaranthaceae
Hectorellaceae
= Portulacaceae
Erycibaceae
Heliotropiaceae
Erythropalaceae
Heloniadaceae
Eucryphiaceae
Helosidaceae
Euryalaceae
Henriqueziaceae
Exocarpaceae
Hippocastanaceae
Flindersiaceae
Hippocrateaceae
Foetidiaceae
Hippuridaceae
= Convolvulaceae
= Olacaceae
= Cunoniaceae
= Nymphaeaceae
= Santalaceae
= Rutaceae
= Lecythidaceae
= Boraginaceae
= Melanthiaceae
= Balanophoraceae
= Rubiaceae
= Sapindaceae
= Celastraceae
= Plantaginaceae
Volume 85, Number4

1998
(cont'd.)
Hortoniaceae
Lepuropetalaceae
Hostaceae
Lilaeaceae
= Monimiaceae
= Agavaceae
Humbertiaceae
= Convolvulaceae
Hydrastidaceae
= Ranunculaceae
= Pamassiaceae
= Juncaginaceae
Limoniaceae
= Plumbaginaceae
Liriodendraceae
= Magnoliaceae
Hydrocotylaceae
Lobeliaceae
Hydropeltidaceae
Lomandraceae
Hydrophyllaceae
Lophiraceae
Hymenocardiaceae
Lophophytaceae
Hypecoaceae
Luxemburgiaceae
Hypericaceae
Malaceae
Hypseocharitaceae
Martyniaceae
Idiospermaceae
Mastixiaceae
Illecebraceae
Medeolaceae
Jasionaceae
Meliosmaceae
= Araliaceae
= Nymphaeaceae
= Boraginaceae
= Euphorbiaceae
= Papaveraceae
= Clusiaceae
= Geraniaceae
= Calycanthaceae
= Caryophyllaceae
= Campanulaceae
Jasminiaceae
= Oleaceae
= Campanulaceae
= Laxmanniaceae
= Ochnaceae
= Balanophoraceae
= Ochnaceae
= Rosaceae
= Pedaliaceae
= Comaceae
= Liliaceae
= Sabiaceae
Mendonciaceae
= Acanthaceae
Johnsoniaceae
Mesembryanthemaceae
Julianiaceae
Mimosaceae
Kiggelariaceae
Monotaceae
Kingdoniaceae
Monotropaceae
Kirengeshomaceae
Mouririaceae
Labiatae
Moutabeaceae
Langsdorffiaceae
Myriophyllaceae
Leeaceae
Mystropetalaceae
Leguminosae
Najadaceae
Leitneriaceae
Nandinaceae
Lemnaceae
Napoleonaceae
Lennoaceae
Naucleaceae
Leoniaceae
Nectaropetalaceae
= Hemerocallidaceae
= Anacardiaceae
= Flacourtiaceae
= Circaeasteraceae
= Hydrangeaceae
= Lamiaceae
= Balanophoraceae
= Vitaceae
= Fabaceae
= Simaroubaceae
= Aizoaceae
= Fabaceae
= Dipterocarpaceae
= Ericaceae
= Memecylaceae
= Polygalaceae
= Haloragaceae
= Balanophoraceae
= Hydrocharitaceae
= Berberidaceae
= Lecythidaceae
= Araceae
= Boraginaceae
= Violaceae
= Rubiaceae
= Erythroxylaceae
551
Annals of the
552
(cont'd.)
Nelsoniaceae
Pistiaceae
Nemacladaceae
Platystemonaceae
Nesogenaceae
Plumeriaceae
Nolanaceae
Podoaceae
Nolinaceae
Podophyllaceae
Nupharaceae
Polygonanthaceae
Nyctanthaceae
Potaliaceae
Nyssaceae
Ptaeroxylaceae
Octoknemaceae
Pteridophyllaceae
Oftiaceae
Punicaceae
Ophiopogonaceae
Pyrolaceae
Osyridaceae
Ranzaniaceae
Pachysandraceae
Reaumuriaceae
Palmae
Retziaceae
Papilionaceae
Rhinanthaceae
Peganaceae
Rhodoleiaceae
Pentastemonaceae
Rhopalocarpaceae
Peperomiaceae
Rhynchothecaceae
Periplocaceae
Roxburghiaceae
Peripterygiaceae
Ruscaceae
Petermanniaceae
Saccifoliaceae
Petiveriaceae
Salaciaceae
Philadelphaceae
Salicorniaceae
Phormiaceae
Salpiglossidaceae
Phylicaceae
Sambucaceae
Picrodendraceae
Samolaceae
Pinguiculaceae
Saniculaceae
Pistaciaceae
Sarcophytaceae
= Acanthaceae
= Campanulaceae
= Cyclocheilaceae
= Solanaceae
= Convallariaceae
= Nymphaeaceae
= Oleaceae
= Comaceae
= Olacaceae
= Scrophulariaceae
= Convallariaceae
= Santalaceae
= Buxaceae
= Arecaceae
= Fabaceae
= Nitrariaceae
= Stemonaceae
= Piperaceae
= Apocynaceae
= Cardiopteridaceae
= Colchicaceae
= Phytolaccaceae
= Hydrangeaceae
= Hemerocallidaceae
= Rhamnaceae
= Euphorbiaceae
= Lentibulariaceae
= Anacardiaceae
= Araceae
= Papaveraceae
= Apocynaceae
= Anacardiaceae
= Berberidaceae
= Anisophylleaceae
= Gentianaceae
= Rutaceae
= Papaveraceae
= Lythraceae
= Ericaceae
= Berberidaceae
= Tamaricaceae
= Stilbaceae
= Orobanchaceae
= Hamamelidaceae
= Sphaerosepalaceae
= Ledocarpaceae
= Stemonaceae
= Convallariaceae
= Gentianaceae
= Celastraceae
= Amaranthaceae
= Solanaceae
= Adoxaceae
= Primulaceae
= Apiaceae
= Balanophoraceae
Volume 85, Number4

1998
(cont'd.)
Sarcospermataceae
= Sapotaceae
Sargentodoxaceae
= Lardizabalaceae
Saurauiaceae
= Actinidiaceae
Sauvagesiaceae
= Ochnaceae
Scaevolaceae
= Goodeniaceae
Scepaceae
= Euphorbiaceae
Schoepfiaceae
= Olacaceae
Sclerophylacaceae
= Solanaceae
Scoliopaceae
= Liliaceae
Scybaliaceae
= Balanophoraceae
Scytopetalaceae
= Lecythidaceae
Selaginaceae
= Scrophulariaceae
Sesamaceae
= Pedaliaceae
Sesuviaceae
= Aizoaceae
Simethidaceae
= Hemerocallidaceae
Siphonodontaceae
= Celastraceae
Sonneratiaceae
= Lythraceae
Spigeliaceae
= Loganiaceae
Stenomeridaceae
= Dioscoreaceae
Sterculiaceae
= Malvaceae
Stilaginaceae
= Euphorbiaceae
Strychnaceae
= Loganiaceae
Stylobasiaceae
= Surianaceae
Stylocerataceae
= Buxaceae
Symphoremataceae
= Lamiaceae
Syringaceae
= Oleaceae
Tetracentraceae
= Trochodendraceae
Tetradiclidaceae
= Peganaceae
Tetragoniaceae
= Aizoaceae
Tetrastylidiaceae
= Olacaceae
Thalassiaceae
= Hydrocharitaceae
Theligonaceae
= Rubiaceae
Thunbergiaceae
= Acanthaceae
Tiliaceae
= Malvaceae
Trapaceae
= Lythraceae
Trapellaceae
= Pedaliaceae
Tribulaceae
= Zygophyllaceae
Tricyrtidaceae
= Liliaceae
Trilliaceae
= Melanthiaceae
Triplostegiaceae
= Valerianaceae
Uapacaceae
= Euphorbiaceae
Ullucaceae
= Basellaceae
Umbelliferae
= Apiaceae
Utriculariaceae
= Lentibulariaceae
Uvulariaceae
= Colchicaceae
Vacciniaceae
= Ericaceae
Viburnaceae
= Adoxaceae
Viscaceae
= Santalaceae
Viticaceae
= Lamiaceae
Walleriaceae
= Tecophilaeaceae
Wellstediaceae
= Boraginaceae
Xanthophyllaceae
= Polygalaceae
Xerophyllaceae
= Melanthiaceae
Zannichelliaceae
= Potamogetonaceae
553

(APG, 1998) An Ordinal Classification For The Families of Flowering Plants (APG I)

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(APG, 1998) An Ordinal Classification For The Families of Flowering Plants (APG I)

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An Ordinal Classification for the Families of Flowering Plants

Author(s): The Angiosperm Phylogeny Group

The AngiospermPhylogeny Group'

Why rearrangefamilies,still less formalizeorders? Higher-levelclassifications,

ANN. MISSOURI BOT. GARD. 85: 531-553.

We note thatthe selectionof clades to be repand in positioninggeneraof

Volume 85, Number4

latterconsistsofa singlefamily.Examplesare Ca- thatconceivedbyBremeret al. (1995, 1996, 1997)

D. H. Les, B. D. Mishler,M. R. Duvall, R. A. Price,

theybelongneitherin any of the phylogenetically

Volume 85, Number4

matK and rbcLsequence data. Amer.J. Bot. 84: 565580.

CLASSIFICATION OF FLOWERING PLANTS

Volume 85, Number4

CLASSIFICATION OF FLOWERING PLANTS

CLASSIFICATION OF FLOWERING PLANTS

Volume 85, Number4

CLASSIFICATION OF FLOWERING PLANTS

CLASSIFICATION OF FLOWERING PLANTS

Volume 85, Number4

CLASSIFICATION OF FLOWERING PLANTS

CLASSIFICATION OF FLOWERING PLANTS

Buxales Takht.ex Reveal

A. C. Sm. ex J. -F. Leroy

Volume 85, Number4

CLASSIFICATION OF FLOWERING PLANTS

CLASSIFICATION OF FLOWERING PLANTS

- not accepted, family

Volume 85, Number4

CLASSIFICATION OF FLOWERING PLANTS

CLASSIFICATION OF FLOWERING PLANTS

TofieldialesReveal & Zomlefer

VellozialesR. Dahlgrenex Reveal

Volume 85, Number4

CLASSIFICATION OF FLOWERING PLANTS

CLASSIFICATION OF FLOWERING PLANTS

Volume 85, Number4

CLASSIFICATION OF FLOWERING PLANTS

CLASSIFICATION OF FLOWERING PLANTS

Volume 85, Number4

CLASSIFICATION OF FLOWERING PLANTS

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