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Systematic Botany 25 Phylogenetic Relationships in the Commelinaceae: I. A. Cladistic Analysis of Morphological Data Author(s): Timothy M. Evans, Robert B. Faden, Michael G. Simpson, Kenneth J. Sytsma Source: Systematic Botany, Vol. 25, No. 4 (Oct. - Dec., 2000), pp. 668-691 Published by: American Society of Plant Taxonomists Stable URL: http://www.jstor.org/stable/2666727 Accessed: 10/09/2010 13:35 Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at http://www.jstor.org/page/info/about/policies/terms.jsp. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at http://www.jstor.org/action/showPublisher?publisherCode=aspt. Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed page of such transmission. JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. American Society of Plant Taxonomists is collaborating with JSTOR to digitize, preserve and extend access to Systematic Botany. http://www.jstor.org SystemnaticBotany (2000), 25(4): pp. 668-691 ? Copyright2000 by the AmericanSociety of Plant Taxonomists PhylogeneticRelationships in the Commelinaceae: I. A Cladistic Analysis of Morphological Data TIMOTHY M. EVANS1 Departmentof Botany,University of Wisconsin,430 LincolnDrive, Madison, Wisconsin 53706 Presentaddress, author for correspondence: Department of Biology,Hope College,35 East 12thStreet, Holland,Michigan 49423-9000 ROBERT B. FADEN Departmentof Botany,NHB 166,National Museum of NaturalHistory, Smithsonian Institution, Washington,DC 20560 MICHAEL G. SIMPSON Departmentof Biology,San Diego StateUniversity, San Diego, California92182 KENNETHJ. SYTSMA Departmentof Botany,University of Wisconsin,430 LincolnDrive, Madison, Wisconsin 53706 CommunicatingEditor: Richard Jensen ABSTRACT. The plantfamily Commelinaceae displays a wide rangeof variation in vegetative,floral, and inflorescencemorphology. This high degree of variation,particularly among charactersoperating under strongand similarselective pressures (i.e., flowers), has made the assessmentof homology among morpho- logical charactersdifficult, and has resultedin severaldiscordant classification schemes for the family. Phy- logeneticrelationships among 40 of the 41 generain the familywere evaluatedusing cladisticanalyses of morphologicaldata. The resultingphylogeny shows some similarityto themost recent classification, but with some notabledifferences. Cartonemna (subfamily Cartonematoideae) was placed basal to therest of the family. Triceratella(subfamily Cartonematoideae), however, was placed amonggenera within tribe Tradescantieae of subfamilyCommelinoideae. Likewise, the circumscriptionsof tribesCommelineae and Tradescantieaewere in disagreementwith the mostrecent classification. The discordancebetween the phylogenyand the most recentclassification is attributedto a high degreeof convergencein variousmorphological characters, par- ticularlythose relating to the androeciumand the inflorescence.Anatomical characters (i.e., stomatal struc- ture),on theother hand, show promisefor resolving phylogenetic relationships within the Commelinaceae, based upon theiragreement with the most recent classification. The Commelinaceae,a well definedfamily of 41 neae is foundmainly in Africaand Asia. Only six genera and about 650 species,is characterizedby genera (Aneilema,Buforrestia, Commelina, Floscopa, several featuresincluding a distinctclosed leaf Murdannia,and Pollia)have indigenousspecies in sheath,a succulentleaf blade, and three-merous bothhemispheres. flowerswith distinctpetals and sepals (Cronquist Older classificationsof the Commelinaceae relied 1981; Faden 1985;Faden and Hunt 1991).The gen- heavilyon floralfeatures (see Fadenand Hunt1991, era are largelytropical and subtropical,but several fora review).Woodson (1942) firstused inflores- extendinto temperateregions. The greatestdiver- cence charactersfor higher-level classification, but sity is in Africa,where, along with Madagascar, he was largelyunfamiliar with the Old Worldgen- nearlyhalf the genera and about40% ofthe species era. Pichon(1946) employedanatomical characters are found(Faden 1983a). to separatethe genus Cartonemaas the familyCar- There is a naturaldivision of taxa betweenthe tonemataceae,but he returnedto more traditional Old and the New World.The seven subtribesof charactersto definehis ten tribesof Commelina- tribeTradescantieae (sensu Faden and Hunt 1991) ceae. Brenan(1966) used a varietyof morphological are each whollyconfined to eitherthe Easternor charactersto define15 informal"groups" of gen- Westernhemisphere, whereas the tribeCommeli- era. These have served as the basis of surveysof 668 2000] EVANS ET AL.: COMMELINACEAE 669 anatomical(Tomlinson 1966, 1969) and cytological molecularand morphologicalstudies are similarto (Jonesand Jopling1972) charactersin the family, the molecularphylogenies (e.g., Chase et al. 1995; but theyhave littletaxonomic significance. Linderand Kellogg1995). Recent work by Givnish, The mostrecent classification, which will be used Evans,Pires, and Sytsma(Givnish et al. 1995,1999) in thisstudy, was put forwardby Faden and Hunt generallysupports these findings. It also places the (1991).In theirclassification, the Commelinaceaeis Hanguanaceae as the sisterfamily of the Comme- divided into two unequal subfamilies(Table 1). linaceae and the Xyridaceaeand Eriocaulaceaein SubfamilyCartonematoideae contains the tribes the grass/sedgeclade. Cartonemateaeand Triceratelleae,each comprising Certainmorphological characters also lend sup- a single genus. SubfamilyCommelinoideae con- port to the separationof the Commelinaceaefrom tainsthe remaining38 generathat are arrangedin otherfamilies of the originalCommelinales (sensu two tribes,Tradescantieae and Commelineae,the Cronquist1981, or Dahlgrenet al. 1985). As dis- formerof whichis dividedinto seven subtribes. cussed above,Dahlgren et al. (1985) separatedthe Evans (1995)conducted a cladisticanalysis of the Commelinaceaefrom the othercommelinoid fami- Commelinaceaeusing both morphological and mo- lies in partby the presenceof raphides,which are lecularcharacters. This paper representsa modifi- absentin theMayacaceae, Rapateaceae, Xyridaceae, cation and expansionof the morphologicalcom- and Eriocaulaceae (Dahlgren and Clifford1982). ponentsof thatstudy. Raphides are widespread,however, in the Ponted- Relationshipsof Commelinaceaeto otherMono- eriaceae,Philydraceae, and Haemodoraceae(Dahl- cot Families. The familyCommelinaceae is the gren et al. 1985). Additionally,whereas the Com- namesake forthe order Commelinales.Cronquist melinaceaewere separatedfrom other families in (1981) definedthe orderby thepresence of perfect the Commelinalesby thepresence of an amoeboid flowersthat are adapted for generalinsect polli- tapetum,the genus Pontederiaand all investigated nation,having showy petals thatare differentiatedmembers of the Haemodoraceae also have an amoe- fromthe sepals. He includedthree other families in boid tapetum(Dahlgren and Clifford1982; Simp- this order:Xyridaceae, Rapateaceae, and Mayaca- son 1988, 1990). Finally,flowers in the Commeli- ceae. He separated the Commelinaceaefrom the naceae show a strongtendency toward zygomor- other three families by their well-definedand phy,whereas they are generallyactinomorphic in closed leaf sheath,and thesucculent blade. the other membersof the Commelinalessensu Dahlgrenet al. (1985)included the same families Cronquist(1981) or Dahlgrenet al. (1985). Zygo- in Commelinalesas Cronquist,but with the addi- morphicflowers are common,however, in thePon- tionof theEriocaulaceae. Eriocaulaceae was placed tederiaceae,Philydraceae, and Haemodoraceae. in itsown orderby Cronquist,who maintainedthat Althoughnumerous taxonomic treatments have it was mostlikely derived within Xyridaceae. Dahl- been produced forthe Commelinaceae,there has grenet al. (1985) distinguishedthe Commelinaceae been littleconsensus as to whichcharacters should fromthe othercommelinoid families by the pres- be used to definerelationships among the genera. ence of raphides,an amoeboid tapetum,and the The earliersystems relied heavily upon featuresof leaf charactersused by Cronquist (closed leaf the highlyvariable androecium, but theyignored sheathand succulentblade). charactersof the inflorescence.The classification Although cladistic analyses of morphological producedby Faden and Hunt (1991),which will be data supporta monophyleticCommelinales sensu used throughoutthis study,incorporated a broad Dahlgrenet al. (1985;Stevenson and Loconte1995), rangeof information,such as charactersfrom mor- recentlypublished moleculardata suggest other- phology,anatomy, and palynology.None of these wise. Nucleotidesequence data fromthe chloroplast classifications,however, is rootedin an evolutionary gene rbcL(Chase et al. 1993;Clark et al. 1993;Du- framework.The purpose of this studywas to ex- vall et al. 1993) place the Commelinaceaenear
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