Delivered by Publishing Technology to: University of Missouri-Columbia IP: 128.206.9.138 on: Wed, 17 Aug 2011 13:14:52 Copyright (c) American Society for Plant Taxonomists. All rights reserved. © Copyright2010bythe American SocietyofPlantTaxonomists Systematic Botany described asmany29speciesof Coniandrae ( Jeffrey 2005 ). Historically, taxonomicworkshave the tribeGuraniinae( Jeffrey 1963 ), butare nowembeddedin Cogn. ( Kocyan etal.2007 ). Thethree were onceclassifiedas genus ismostcloselyrelated to ring indryandwetforests atelevationsupto2,100m.The much more oftenthanfemaleflowers; therefore, theirchar- culate racemes orspikes.Maleflowersare seeninnature tudinal splotches. on somespecieshavesmall, darker green specklesor longi- Vegetative structures are brightgreen, and stemsandtendrils emerge from thestem ata90degree anglefrom theleafnode. 1916 ; Hampshire 1992 ). Thevineshavesimpletendrils,which they were usedintaxonomickeysto used todistinguishspeciesconfidently, althoughhistorically cies. Thisisonereason whyleafcharactersalonecannotbe variation isinconsistentamongindividualsofthesamespe- lowly trilobedleafadjacenttoonethatistrifoliolate,andthis the sameindividual.Itisnotuncommontoseeasimple,shal- ferent leafshapesoccuronthevariousspecies,andeven along theveinsandmargins ofsomespecies. A numberofdif- are typicallyglabrous, butminute,softtrichomesmaybeseen and canreach lengthsof75mmandwidths53mm.Leaves leathery. Theleavesare arrangedalternatelyalongthestem, some speciesbecomewoody, andsome mature leavescanbe herbaceous withmembranousleaves,whileolderplantsof stems andleavesofotherplants. Young tops ofshrubs, climbingwithtendrilsthatwraparound the at thatlevel.Theplantsare typicallyseengrowing overthe canopy, butitisunknownhowextensive the forest floor(pers.obs.). Thevinescangrow upintothe tances betweenindividualsofonekmormore atthelevelof 2010). lar phylogenyof here basedonmorphological studies,andarecent molecu- ers haveledtoaninflatednumber. Sixspeciesare recognized system consistingoftemporallyandspatiallyseparatedflow- over thelifeofanindividualandamonoeciousreproductive characteristics suchasleafandflowermorphologythatvaries Psiguria From someleafnodes,maleflowersemerge onlongpedun- Psiguria Section ofIntegrativeBiology, TheUniversityofTexas at Austin, 1UniversityStationC0930, Austin, Texas 78712U.S. A. identify speciesof list ofcharactersthatdistinguishallpairsgeographicallyoverlappingspecies,botanistsandecologistsfinallyhavethe another thatincorporatesleafcharacteristicsandgeographicdistributionbutuniquelyidentifiesspeciesaccording toDNA bar sented. A descriptionandadistributionmapofeachspeciesisprovided alongwithtwotaxonomickeys:onethatutilizesmale six species.Inthisrevision, areview ofover35yrpublicationsaddressing ecologicalandnaturalhistorystudiesfocusin mens andindividualsfrom bothgreenhouse andnaturalsettings,combinedwitharecent molecularphylogeny, supportsthedeline by misleadingcharacteristicssuchasvariableleafandflowermorphology. A thorough morphologicalinvestigationof758herbar fly andhummingbird pollinators.Historically, taxonomicrevisions havedescribedasmany29species, butthisnumberhasbe and shade. Keywords— Abstract— Arn. isagenusofperennial, Neotropical vinesoccur- israre throughout itsgeographicalrangewithdis- (2010), 35(2):pp.341–357 Psiguria aooi Rvso o te etoia Gns Genus Neotropical the of Revision Taxonomic Psiguria Psiguria N broig, barcoding , DNA Psiguria anditssistergenera, isagenusoflianasfoundthroughout theNeotropics from sealevelto2,100m,indryorwetforests, inbothlightgaps Current address: UniversityofMissouri-Columbia, 1201RollinsSt.-BondLSC311, supportedthisnumber(Steeleetal. . Guraniinae Gurania Columbia, Missouri65211 U.S. A. ( [email protected] ) Psiguria Psiguria Gurania Cogn.and , Heliconius omnctn Eio: nra Schwarzbach Andrea Editor: Communicating Psiguria Psiguria , butmisleading and (e.g. Cogniaux Helmontia , Helmontia growth is vinesare Helmontia P Rxne Steele Roxanne P. , are uniqueamongCucurbitaceaehavingbrightlycolored flowersandbutter- rpcl vines. tropical ,

341 batus species, currently placedin been usedforidentifyingspecies of nature, theirmorphologicalcharacteristicshavenottypically and whitestriped.Becausefemale flowersare rarely seenin hiscent, cucumber-like fruit thatmaybedarkgreen orgreen fertilization, theinferiorovary developsintoasmall,inde- just beseenemerging from thetubeofamature flower. Upon two. Thestigmaticsurfacesare rugose, andtheirendsmay divided intotwo,andthesesectionsare furtherdividedinto and atanode.Flowersare epigynouswithastigmathatis At agiventime,manyfemaleflowersmaybeopenonplant vine thatmaybeleaflessorcontinuetoproduce mature leaves. appendage isinconsistent. of manyspecimensrevealed thatthepubescenceofanther of rates guish species.Whilefoldedanthershapedefinitivelysepa- throat were originallyusedby Cogniaux (1916) todistin- and thepubescenceofsmallantherappendagesneartube inner floraltube.Theshapeoftheanthers(straightorfolded) between flowersonthesameinflorescence. revealed thatthischaracteristic isinconsistentandmayvary cylindrical orflask-shaped;however, widespread sampling used by Wunderlin (1978) wastheshapeofcalyxtube, tent, are includedinthepresent keys. Another characteristic Wunderlin (1978) and,becausetheywere foundtobeconsis- lengths mayvary. Thesetwocharacteristicswere usedby species havedarkergreen speckles,andsepalshapes various shadesofpink,red, ororange.Thecalicesofsome lar green calyxtubeand outspreading petals.Petalsmaybe others havepedicelsupto19mmlong. millimeter. Additionally, somespeciesare apedicellatewhile together thatmore thanone flowerscarmaybeobservedper the lengthofpedicels.Insomespecies,flowersoccursoclose the numberofflowerspermillimeteralongpeduncleand characteristics thathavebeenoverlookedinprevious keysare acteristics are bestused to identifyspeciesof Nmnltrl History— Nomenclatural Female flowerstypicallyemerge atnodesontheapexof Inside themaleflowersare onlytwoanthersattachedtothe Both maleandfemaleflowersare salverform,withatubu- Psiguria L., Psiguria pedata C. trifoliata , inwhichtheanthersare straight,theexamination Psiguria L.,and (L.)R. A. Howard from allotherspecies (Cucurbitaceae) Lnau (79 ecie three described (1759) Linnaeus C. pedatus Psiguria Psiguria , under L. Von Jacquin(1760) g on tools toconfidently . Psiguria codes. With a flowers, and Cucumis trilo- Psiguria en inflated ium speci- ation of ispre- . Two . Delivered by Publishing Technology to: University of Missouri-Columbia IP: 128.206.9.138 on: Wed, 17 Aug 2011 13:14:52 Copyright (c) American Society for Plant Taxonomists. All rights reserved. P. ovata ottoniana Nonetheless, Jeffrey (1978) madesixnewcombinations; too manyspecies,andincludednumerous misidentifications. also argued thatCogniaux, inhis1916treatment, recognized far Anguria Psiguria Wunderlin. Inthesameyear, Jeffrey (1978) circumscribedpedunculata The relationships between thethree generainthe clade ognized here. rosa, P. warscewiczii,P. triphylla from Steeleetal.2010).Thesesix, support foronlysixspecies of for themonophylyofgenus, aswellweaktostrong ( Schaefer etal.2009 ). Steeleetal.(2010)found strong support Gurania et al.2007 ). Thespeciationeventseparating C. Jeffrey, anddescribedonenew species, P. warmingiana binations; becoming thetype.In1978,Wunderlin madethree more com- P. jacquiniana for tions in be transferred to Jeffrey’s (1962) assertionthatmany and Shaw1966 ; Dandy 1967 that indicatedtheequivalenceof ), Howard (1973) corroborated nal genericdiagnosiswasderivedfrom because von Jacquin’s(1763) descriptionofthefruit inhisorigi- taxonomic scrutiny. Additionally, Jeffrey (1962) explainedthat, but onlyif Cogniaux’s (1916) splitintotwogenerastoodupto Gurania the name gitimate, but Jeffrey (1962) , inhisdiscussionofthehistory species), basedonmorphologicaldifferences. in Subsequently, fouradditionalcombinations were made [Volume 35 SYSTEMATIC BOTANY species underthename Mill. ( Miller 1754 ). moved theseto Linnaeus (1762) publishedthesethree 342 monophyly ofacladeincluding placed and FernandezCasas1998 ). sifolia P. aurantiaca P. dunlapii divided intotwogenera, Anguria 1881). 1876, Cogniaux In 1916,themostrecent andcompletetaxonomicrevision of 1851; Schlechtendal von 1846; ( Roemer several additionalspecieswere describedunder (1762) had usedthename not listanybinomials.Despitethis,andbecause Linnaeus et al.2006 ). Arnott (1841) published (ICBN) hassuppressed allnamesinthatpublication( McNeill however, theInternationalCodeofBotanicalNomenclature and hefirstpublishedthename that thisgenericnamewasincorrectly appliedtothesetaxa, Comparative Morphology andGeographic Distribution— Rickett andStafleu(1960) firstnotedthat Phylogeny— Psiguria Anguria (Cogn.)C.Nelson&Fern.Casas( Nelson Sutherland (Donn.Sm.)C.Jeffrey, isestimatedtobesix ±three millionyears ago , butquestionedthemonophylyof (appliedtothosespeciespreviously describedunder ) and waspublishedbyCogniauxinwhichthegroup was Psiguria Psiguria (Schltdl.)C.Jeffrey, P. bignoniacea (Standley)R.J.Hampshire ( Hampshire 1992 ), Anguria asfollows: shouldbe (Cogn.)Wunderlin, and (S.F. Blake)C.Nelson&Fern.Casas,and (Schltdl.)R. A. Howard withthefirstnamedefacto Gurania (Cogn.)C.Jeffrey, and The mostrecent phylogenyofCucurbitaceae intribeConiandreae, andsupportedthe ; ; Anguria P. pedata,P. trilobata Psiguria , proposed conservingthename basedonfloralsyndromes. Jeffrey (1978) A. pedata (Poepp.&Endl.)Wunderlin, P. trifoliata Jacq.,alaterhomonymof Anguria Anguria , andhemadethree newcombina- Anguria . Along withotherpublications P. umbrosa , and Psiguria P. triphylla Psiguria Psiguria (29species)and . DeNecker(1790)realized (L.) Alain ( Liogier 1980 ), P. pedata,P. ternata,P. umb- , inthemid tolate-1800s, Anguria Psiguria Psiguria P. warscewiczii P. racemosa (L.)R. A. Howard, and A. pedata P. ternata in (Fg 1 reproduced 1, (Fig. with P. racemosa (Kunth)C.Jeffrey, speciesneededto (Miq.)C.Jeffrey, Elementa Botanica , , Psiguria Anguria Arn., buthedid Helmontia Anguria , thelectotype , are alsorec- Psiguria M Roem.) (M. Gurania Ho. f.) (Hook. . Jeffrey. C. (Kocyan wasille- Psiguria Anguria Anguria P. longi- P. diver- (Willis , and , and (73 .P. , ;

brightly colored flowers in genera consistofNeotropical vineswithlarge leaves,and in tetrads( Jeffrey 1964 ; Roubik andMoreno 1991 ). All three Gurania molecular phylogenyof ( have single-grainpollen, sister because,althoughmostcucurbitsincluding len morphologywouldsuggestthat red, pink,ororange, Psiguria but itdoesnotexplainthesimilaritiesbetween Anguria of evolutionary histories.However, themolecularseparation The nucleardatasetspresent conflictingandpoorlysupported house locationssincetheearly1970s( Gilbert 1972 , 1975 , 1977 tory of ; system, ecologicalinteractions,phenology, andnaturalhis- South America. petal color),but Helmontia Paraguay, butonly geographic distributionsstretching from southernMexicoto that are oftenmistakenfor anthers. long, dividedsepalswithshort,inconspicuous,yellowpetals ers havepredominantly pubescent leavesandbrightorange, out beyondthethroat of the tube.Incontrast, short, green sepals,andred, pink,ororangepetalsspreading forms atube(see Fig. 2 forshapevariations)endinginfive brous leavesandsalverform flowersinwhichthegreen calyx Psiguria them from mostothercucurbits ( Condon andSteck1997 ). intron. Bootstrapvaluesare indicatedabovebranches. intergenic spacers,ITS,andthenuclear Steele etal.2010, Fig. 5 ) inferred from acombinationofeightchloroplast drical withthickened base,E.flask-shaped. Psiguria, Gurania Fig. Eooy n Ntrl History— Natural and Ecology Fig. Psiguria 2 1 . . Psiguria . . and flowers thatdiffer mainlyinpetalcolor( basedonstrikingdifferences inflowermorphology, isdistinguishedfrom Floral tube shapes. A. cylindrical, B.elliptical,C.oval,D.cylin- flowersare similartothoseof Pyoey f of Phylogeny and Helmontia havebeenstudiedat severalfieldandgreen- Gurania Helmontia , and Psiguria formacladesisterto Helmontia: Psiguria supports Cogniaux’s (1916) splitof Helmontia Psiguria isknownonlyfrom northeastern Psiguria extendsintotheGreater Antilles. Psiguria (Cucurbitaceae)(reproduced from Gurania wie. diinly pol- Additionally, white). serine/threonine phosphatase basedonplastidgenes, and ) are stillunclear. Inthe Aspects ofthepollination Psiguria and

Psiguria bypredominantly gla- Gurania Gurania Psiguria and and Psiguria Helmontia Gurania shedpollen Gurania distinguish (otherthan Gurania Helmontia Psiguria (Fg 1). (Fig. flow- have gene and are :

Delivered by Publishing Technology to: University of Missouri-Columbia IP: 128.206.9.138 on: Wed, 17 Aug 2011 13:14:52 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 1986 ; Murawski 1987 ; 32:1) hasbeen observed ( Gilbert 1975 Condon andGilbert1988 ; ). Extensive Murawski andGilbert For allspecies,astrongly male-biasedfloweringratio (e.g. peaks oftenatthestartof rainyseason( Condon 1984 ). Additionally, seasonalfluctuationshavebeenobserved,with Murawski andGilbert1986 ; per plantvariesovertime, from zeroMurawski 1987 to46( ; Gilbert 1977 Gilbert 1993 ; but theaveragedailynumber ofinflorescences produced ). 1987 ; Condon andGilbert1990 ). and neitherhasbeendescribedashavingscent( Murawski in thecanopyorlightgapsshadeatlowerlevels, Gilbert 1988 ). Bothmaleandfemaleflowersmayemerge high ing intofruit ( Condon 1984 ; only onceforfewtoseveralhoursbefore fallingoff ormatur- Murawski 1987 ; Condon and may beopenatatime;howeverbothtypesofflowers cence eachday, whileonetoseveralfemale flowersatanode states ( Condon andGilbert1990 ). large vinescanalternatebetweenfloweringandvegetative and fruiting untilitreached 15m in length. Additionally, an eight-yearperiod,alternatedbetweenfemaleflowering La SelvaBiologicalResearch StationinCostaRicathat,over triphylla Condon andGilbert1988 ). Gilbert (1983) observeda continue foranundeterminedamountoftime( Condon 1984 ; to tendaysunlessnofruit isset,inwhichcasefloweringmay Female branchesproduce receptive flowersforaboutaweek to three weeksafterpollination ( Condon andGilbert1990 ). female flowerspernode,withfruits maturingabouttwo Gilbert 1990 ). A plantmayhavefrom onetoasmanysix as twometersbefore producing thefirstflower( Condon and main vine,andtypicallyturndownward, dropping asmuch and Gilbert1990 ). pollinated specieshaveproduced viableoffspring ( Condon may beanadaptationforoutcrossing sinceartificiallyself- 1983 ). Thetemporalseparationofmaleandfemaleflowers seed germinationtofruit setcanbeseveralyears( Gilbert are produced, butthisisrare ( Murawski 1987 ). Thetimefrom plants continuetoproduce maleflowerswhenfemale 343 drils ( Murawski 1987 ; Condon andGilbert1988 , minal branches,sometimeswithreduced leavesandten- 1990 ). Some plants switchsextoproduce femaleflowersatnodesonter- diate flowerlessperiodlastingafewdaystoseveralmonths, 1988 ). Oncestemsreach adequatesize,andafteraninterme- a year( Gilbert 1975 , 1983 ; climbing vines,andproduce flowersforthree months toover Condon 1984 ; Condon andGilbert male inflorescences are firsttoemerge from leafnodesof Condon 1984 ; Condon andGilbert1988 ). Indeterminate,erect spatially separatedmaleandfemaleflowers( Gilbert 1983 ; these plantswere indeedmonoeciouswithtemporallyand until severallong-term,detailedobservationsrevealed that STEELE:TAXONOMIC REVISIONOFPSIGURIA rarely monoecious,members.Thismisdirected idea persisted Cogniaux (1916) describedthegenusashavingdioecious, ( Linnaeus 1759 , him described 1762 ; to bedioecious.Despitethefactthatseveralbotanistsbefore von Jacquin 1760 ; de Necker1790 these investigations,mostspeciesof ), 1997 ; Condon etal.2008 ). From thetimeofCogniauxuntil Gilbert 1993 ; Condon andNorrbom1994 ; Gilbert 1986 ; Condon andSteck Murawski 1987 ; Condon andGilbert1988 , 1990 Boggs etal.1981 ; ; Gilbert 1983 ; Condon 1984 ; Murawski and 2010] Ms Most Zero tothree maleflowersare typicallyopenperinflores- Female branches hangonothervegetationawayfrom the Psiguria (called Anguria speciescanproduce maleflowersyear-round, Anguria pachyphylla taxaunderthecategoryofmonoecious Donn.Sm.)individualat Psiguria were thought Psiguria tors. Although tors. Although the most-studiedofwhichisrelationship withitspollina- and Gilbert1990 ; Gilbert 1993 ). 1977 ; Boggs etal.1981 pollinators, ; Murawski andGilbert1986 ; Condon sources, mayhaveinfluencedspeciationin flowering time,providing nearlyconstantpollenandnectar (along withsistergenus ( Condon andGilbert1990 ). Thisinteractionmakes visit femaleflowers“accidentally”whileforagingforpollen matic surfaces,itishypothesizedthatthebutterfliesmay female flowers,andthesimilartextures ofanthersandstig- Because ofthemorphologicalsimilaritybetweenmaleand 1986 ; Murawski 1987 ; Condon andGilbert1990 ; et al.1981 Gilbert 1993 ). ; Gilbert 1983 ; Condon 1984 stigmas offemaleflowers( ; Murawski andGilbert Gilbert 1972 , visits andtransferringthegreatest amountofpollentothe 1975 , 1977 Heliconius ; Boggs isms suchashummingbirds, bees,moths,andbutterflies, composition ( Murawski andGilbert1986 ; Murawski 1987 ). ers, geneflowintheplantpopulationwillvarywithvisitor isms thatrob pollenrather thantransferringittofemaleflow- However, because genera thatare butterfly orbird-pollinated ( Gilbert 1983 ). (Tephritidae) ( Condon 1984 nitidalis ; Condon andGilbert1990 are associatedwith ; 1975 , 1983 ; Condon andGilbert1990 ). Severalinsect parasites keys destroy theseedsorprey onimmature fruits ( Gilbert Condon andGilbert1990 ). Parrots, squirrels, mice, andmon- primary fruit dispersalagents( Gilbert 1983 ; have beenobservedorstudied. Batsare believedto be the Condon 1984 ; 1987). Murawski of thelong-lived(uptosixmonths)butterflies( Gilbert 1977 ; source contributestothenearlyconstantegg-layingsuccess tion vectorforthewidely-spacedplants,butsteadyfood Gilbert 1993 ). Notonlydoesthissystemprovide apollina- Condon 1984 ; Murawski andGilbert1986 ing bothsexesnearbyfortimesofbreeding ( ; Gilbert 1975 Murawski 1987 ; ; value, whilemalesfeedlaterinthemorningonnectar, keep- feed earlyinthemorning,gatheringpollenforitsnutritional acids from thepollenintotheirsystem.Femalebutterflies ture withtheirproboscis. Thebutterfliesthenabsorbamino anthers, combineitwithnectar, andthenmassagethemix- nectar to Murawski andGilbert1986 ). butterfly’s role astheprimary pollenvector( Condon 1984 ; sufficient portionistransferred tofemaleflowersfulfillingthe duced by ents obtainedfrom the pollen of most speciesof the benefittopollinatorcannotbeunderstated.Infact, 1981 ). Although mostofthe the large sizeofthepollen tetrads( Gilbert 1975 ; Boggs etal. Psiguria Gurania acids andproteins obtained from pollenof butterflies asadults(bothmaleandfemale)assimilateamino enous compoundsnecessaryforeggproduction, other lepidopteransthat,duringlarvalfeeding,obtainnitrog- Psiguria Heliconius Several otherplant-animalinteractionsinvolving Stoll(Pyralidae),and fruit flies, species( Gilbert 1972 , 1975 , 1977 ; Boggs etal.1981 ). and Heliconius butterflieshavebeenobservedmakingthemost playsarole inasuiteofplant-animalinteractions, Psiguria butterfliesprovide avitalservicefor Heliconius Gurania Psiguria Heliconius isactuallydestroyed bythebutterflies,a Psiguria feeders.Butterfliesscrapepollenfrom the Psiguria are preferred overothersources dueto Klukbutterflies( Gilbert 1972 , 1975 , flowersare visitedbyseveralorgan- flowersare visitedbymanyorgan- Gurania butterfliesrely heavilyonnutri- , includingpickleworm, Heliconius Psiguria ) amongthefewcucurbit Psiguria flowersalsoprovide -collected pollenpro- Blepharoneura Psiguria foes Unlike flowers. Psiguria Psiguria andafew Heliconius Diaphania ’ major ’s Psiguria Psiguria Loew , but ,

Delivered by Publishing Technology to: University of Missouri-Columbia IP: 128.206.9.138 on: Wed, 17 Aug 2011 13:14:52 Copyright (c) American Society for Plant Taxonomists. All rights reserved. depth. ium specimens. A speciesisdefinedasthesmallestmonophyleticgroup cal observationsandmeasurements from field,greenhouse, andherbar- phylogenetic results ofSteeleetal.(2010)withthecurrent morphologi- logenetic speciesconceptsare incorporatedbyaligningthemolecular history. Toward thatend,componentsofbothmorphologicalandphy- descriptions andkeys. rosa included 25- by thegeographicandmorphologicalvariabilityofeachspecies.This may notbeobvious.Thenumberofspecimensmeasured wasdetermined diagram oftwomeasurements (leafbasedepthandlobedepth),which light microscope (formeasurements lessthan10mm). Figure 3 showsa measurements greater than10mm),orthereticule ofanOlympusSZ61 were measured usingeitherametricruler withmillimeterdivisions (for individuals from herbariaorlivingmaterial,43quantitativecharacters Texas at Austin were collectedandfixedinFAA before measuring. On178 Several flowersfrom plantsgrowing ingreenhouses atTheUniversityof by soakingthemindiluted Aerosol OTsurfactantforamaximumof24hr. tographs orcopiesoftypedrawings.Preserved flowerswere rehydrated specimens were examinedwhere possible,someofwhichincludedpho- nata parentheses isnumberofcollections): 4 SSEAI OAY [Volume 35 sentatives from eachofthesixspecies GH, andUS.Theexaminedcollectionstotaled758,includedrepre- and specimenswere borrowed from fiveadditionalherbaria;MO,NY, F, SYSTEMATIC BOTANY Personal collectionsare depositedinTEXandcountriesofcollection, lections coveringthegeographicandmorphologicalrangesof cally similar, sympatricspeciesof Most recently ithasbeendiscovered thatseveralmorphologi- Condon 1984 ; on seedscausingpredispersal seedmortality( Murawski 1987 Gilbert 1983 ; ; female flowers,repellingCondon andGilbert1990 butterflypollinators,andfeeding ). et al.2008 ). Thelatterofthesetwoinfestbothmaleand Condon andNorrbom1994 ; Condon andSteck1997 ; Condon 344 rate partsof 1994 ; Condon andSteck1997 ; Condon etal.2008 as 13different). Fig. pce Concept— Species The present treatment isbasedonbothherbariumandpersonalcol- ad 35- and , 7) (73), 3 . . P. triphylla Diagram of two leafmeasurements. a. basedepth,b.lobe P. pedata P. warscewiczii Psiguria Blepharoneura 27, (297), 11- , aeil ad Methods and Materials Species are distinguishedbytheirevolutionary and P. racemosa . Thesedataservedasthebasisforspecies P. umbrosa Gurania species( Condon andNorrbom 19- , (43),and P. pedata plantsthatcanhostasmany P. ternata Blepharoneura Psiguria 7) (77), P. warscewiczii 65- , asfollows(numberin P. racemosa P. triphylla feedonsepa- 20. Type (250). 23- , 1) (18), Psiguria P. umb- P. ter- . species delineation. type specimens.Whenthisisnotpossible,DNA barcodes are provided for based oncharactersobservedinthefieldand/orbycomparingthemwith that, wheneverpossible,researchers canidentifyorganisms usingkeys phorants are notconfusedasdifferent species.Speciesare describedsuch tory, reproductive biology, andphenologyare considered suchthatsema- races whichhavenotbeendistinguishedgenetically, andthenaturalhis- or more morphologicalcharactertraits.Thesecriteriaexcludeformsor having weaktostrong bootstrapsupportandconsistentlysharingone Psiguria surface glabrous orpuberulent, withorwithouttrichomes at or solid,(4.9–)11.0–41.0 mmlong, 0.9–10.0mmwide,adaxial drical orvariousflaskshapes, glabrous orpuberulent, striped ers 2–5pernode,axillaryorterminal; calyxsalverform,cylin- portion (ifapplicable)(0.7–)1.0–1.5 mmlong.Pistillateflow- or triangular, glabrous orpapillose,0.1–1.2 mmlong,folded backwards, (1.0–)3.4–11.0 mmlong,antherappendageround and dorsifixedinsidefloraltube, bilocular, straightorfolded length-to-width ratio1.2–2.1;anthers2,free, afilamentous rounded toacuminate,2.0–62.0mmlong,2.0–40.0wide, venation reticulate, pubescentinternallyandexternally, apex calyx tube,lobes5,free, orange,pink,orred, ovatetoobovate, darker thancalyx,0.5–8.5mmlong;corolla rotate, adnateto sepals 5,thinorthick,triangularlinear, color sameasor glabrous orpuberulent withorwithouttrichomesatthroat; les, 4–17.0(–21.0)mmlong,1.0–7.0wide,adaxialsurface or puberulent, solidgreen orlightgreen withdarkerspeck- ers, receptacle withtrichomes;calyxsalverform,glabrous 2–3 flowerssometimesmuchlarger thansubsequentflow- with 0.24–1.2flowerspermm.Staminateaxillary, first pedicels absentor, ifpresent, thenpubescent,0–19mmlong, splotches, 0.5–4.6(–8.0)mmindiameter, (8–)21–395mmlong; glabrous orpuberulent, solidgreen orlightgreen withdarker Inflorescences pedunculate racemesorcorymbs;peduncles ate andabaxialbasesacute-truncate, 18–115(–145) mmlong. leaflets onpedateleavesoblique,adaxialbasesacute-attenu- oblique, 25–225mmlong,13–124wide;basesofouter bases ofsideleafletsacute,attenuate,truncate, sometimes acute orattenuate,40–305mmlong,17–132(–150)wide; ceolate, apicesacuminateorcuspidate;basesofcenterleaflets let elliptictooblanceolate,sideleafletslanceolateoblan- diameter, 0–48mmlong;compoundleaveswithcenterleaf- glabrous orpuberulent, sometimes winged, 0.6–3.6mmin wide, lobedepthaverage62.5%oftotallength;petiolules deep, apicesacuminate,39–219mmlong,22–122(–178) ulate, truncate, rounded, and/oracute, bases5–33(–40)mm long; simpleleaveslanceolate,elliptic,ortrilobed,baseauric- ioles glabrous orpubescent, 0.5–5.3mmindiameter, 3–96mm blade, simple,trifoliolate,pedate,oracombinationofthese;pet- sometimes withirregularly-spaced veinsextendingbeyond main veinsflushorprominent, margins entire orirregular, veins andmargins sometimes puberulent, venationreticulate, green, lightergreen below, andbothsurfacesglabrous with the bladesmembranousorcoriaceous,uppersurfacedarker darker speckles,0.2–3.1mmindiameter. Leavespetiolate, eter; tendrilsglabrous orpuberulent, solidorlightgreen with light green withdarker, linearsplotches,0.9–7.0mmindiam- age; stems,minutelysulcate,glabrous orpuberulent, solidor ceous butsomebecomingwoodywithgreater stemsizeand Monoecious, perennial, vineswithsimpletendrils,herba- Psiguriapedata Arn., Hook.Jour. Bot.3:247.1841.—LECTOTYPE: aooi Treatment Taxonomic (L.)R. A. Howard. Delivered by Publishing Technology to: University of Missouri-Columbia IP: 128.206.9.138 on: Wed, 17 Aug 2011 13:14:52 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 1 1. 1. Leavessimple,trilobedor5-lobed,sometimesdeeply, butnevertrifoliolate . 1. Anther straight . 345 . 1. Anther folded backwards STEELE:TAXONOMIC REVISIONOFPSIGURIA oblong, 6.5–22.0mmlong,1.9–7.0wide,ovulesnumerous, to-width ratio1.3–3.5;gynoecium1,bilocular, ovaryelliptic- to acuminate,4.5–16.0mmlong,2.4–9.5wide,length- vate, venationreticulate, bothsidespubescent,apexrounded to calyxtube,lobes5,free, orange,pink,orred, elliptic toobo- as ordarkerthancalyx,0.5–3.7mmlong;corolla rotate, adnate throat; sepals5,thinorthick,triangularlinear, colorsame 2010] below refers tomarkersandreference sequencesfrom Table 4inthatpublication. et al.(2010),whichcontainsalistofchloroplast DNA markersfordistinguishingthesixspeciesof stem andleafcharactersplusgeographiclocation,thenutilizeDNA barcodes forexactidentificationasdescribedinSteel identification. Inthiscase,itwillbenecessarytonarrow theidentificationdowntotwoorthree specieswithafewconsist leaf morphologyofa vines intropical forests are oftenseenwithoutflowers,itmaybenecessarytoidentifysterilespecimens.However, becauset 1. Leavesbifid,trifoliolate,orpedate,withimmature leavesoftensimpleandlobed,orsometimes acombinationofshap (sterile orcarpellatespecimens) (specimens withstaminateflowers) Two keysare provided: Key1forspecimenswithstaminateflowers,and2sterileorcarpellatespecimens.Because Psiguria pedata 2 Flowers 2. 2 Flowers 2. 2 hools DA ein region DNA Chloroplast 2. 3. Stemsurfacesolidgreen, leavesbifid,trifoliolate,pedate,withimmature leavesoftensimpleandlobed, regions DNA Chloroplast 2. 3. Stemsurfacelightgreen withdarkergreen splotches, leavestrifoliolate,pedate,orseldomsimpleandtrilobed,larg 441. 1973. fructu I: 22p.10.1876.—TYPE: pedata Anguriapolyphyllos 1759. (holotype!). 3 Pedicels 3. 3 Pedicels 3. 4. Leafmargins with veins extendingbeyondthemargins oftheleaves,chloroplast DNA region 4. Leafmargins seldom withveinsextendingbeyondthemargins oftheleaves . or sometimesacombinationofcomplexities . in ColombiaorVenezuela . but notmatchingregions lobes onoutsideofsideleaflets,andchloroplast DNA region distribution inVenezuela, Brazil,orLesser Antilles . . 5 eas ik spl hn lna; sepal linear; thin, sepal pink; Petals 5. 5. Petalsorange-red; sepalthick,triangular;0.5–1.5mmlong;colordarkerthan calyx, first2–3flowers 4. Calyxandpedunclecolorspeckled/splotched;sepaldarkerthancalyx;petalsorange . . 4. Calyxandpedunclecolorsolid;sepalssameascalyx;petalspink 5. Leavessimpleandelliptic,2-lobed,trilobed,trifoliolate,ormixedcomplexities; mainveininsideleafletsnot 5. Leavestrifoliolate,orrarely pedate,andmainveininsideleafletsstrongly off-center, typicallywithsmalllobes C.Plumier, Plant. Amer. fasc.I.13 vr var. Anguria pedata P. pedata < > or=to0.75permm;pedicelspresent and 0.75permm;pedicelsabsentor not muchlarger thanlaterflowers;geographicdistributionNofequator . ae lwr;gorpi itiuinSo qao later flowers;geographicdistributionSofequator . in Mexico,Central America, orSouth America . . ndhC-trnV, ndhF-rpL32 strongly off-center, typicallywithoutlobesonoutsidebaseofsideleaflets,chloroplast DNA regions geographic distributioninPeru, Brazil,Bolivia,orParaguay . on outsidebaseofsideleaflets,chloroplast DNA region Cucumis pedatus polyphyllos < > 2.0mmlong;petalapexrounded orrounded-acute . or=2.0mmlong;petalapexacuminate . reference sequence,andgeographicdistributioninGreater Antilles orrarely CostaRica. . (L.)R. A. Howard, J. Arnold Arbor. 54: Psiguria Schltdl.,Linnaea24:712. 1851. Jacq. Enum.Pl.Carib.p.31.1760. (Schltdl.)Cogn.,Diagn.Cucurb. rpoB-trnC rpoB-trnC individualcanchangeoveritslifetime,morphologicalcharactersmaynotbeenoughforaccurate rpoB-trnC L.,Syst.Nat. ed.10p.1279. , and/or Anguria polyphyllos,parvo matches and/or and/or 3rps16-trnQ < > 0.3mmlong . P. racemosa psbZ-trnM 1.5 mmlong;sepalcolorsameascalyx,first2–3flowersmuchlarger than psbZ-trnM tabula 23 Key Key > Key toSpeciesof match 0.5mmlong . match reference sequenceandgeographicdistribution Anguria with 2 1 . 1755. . toSpeciesof toSpeciesof P. triphylla P. umbrosa P. umbrosa ndhF-rpL32 3rps16-trnQ . reference sequences,andgeographicdistribution long, 2.8–5.9mmwide,1.5–3.8thick. tened, withoutadistinctmargin, whitish-gray, 5.0–10.0mm wide, wall0.7–3.0mmthick.Seeds30–88,ovate-oblong,flat- lighter green orwhitestripes,18–80mmlong,10–24 long. Fruit apepo,oblong,indehiscent,solidgreen orwith stigmatic surfacespapillose,stigma3.5–10.2(–12.0)mm horizontal, style1,2.3–9.0mmlong,stigmastwicebranched, Cucumistrilobatus Anguriatrifoliata reference sequencesandgeographic reference 6. sequences . matches P P P SIGURIA lobata eriana (L.) Alain, Phytologia47(3):192.1980.—TYPE: Plumier, Plant. Amer. fasc.I.13 C. 1973.—TYPE: J. ArnoldArbor.442. Howard, 54(4): R. A...... 1. . SIGURIA SIGURIA matches L.,Enum.Syst.Pl.pp.9&31.1760. Schltdl.,Linnaea24:709.1851. P. umbrosa

...... 2. . P. ternata L.Sp.Pl.2p.1376.1763. L., Syst.Nat.ed.10p.1279.1759. eeec sequence, reference ...... 4 . 5. . 2 . reference sequence,and ndhC-trnV 3...... 4. . 4 ...... 2...... 5 . 3. . matches 5 . tabula 22 ...... 6. . Psiguria s ...... 3 es . 5. . e Psiguria trilobata 4. . . 1755.(holotype!)...... 1. . . Thesecondkey Psiguria trifoliata Anguria plumi- P. warscewiczii P. warscewiczii Anguria tri- ...... 3 . P. racemosa P. racemosa P. triphylla P. triphylla P. umbrosa P. umbrosa Cucumis P. ternata P. ternata P. pedata P. pedata (L.) (L.) ent he 2 e

Delivered by Publishing Technology to: University of Missouri-Columbia IP: 128.206.9.138 on: Wed, 17 Aug 2011 13:14:52 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 4 SSEAI OAY [Volume 35 SYSTEMATIC BOTANY Anguriadentata 346 Anguriakeithii Anguriaottoniana 6.3–6.5 mmlong, 3.9–4.1mmwide,2.0–2.4 thick. long, 10–18mm wide,wall0.7–1.3mmthick. Seeds44–46, mm long.Fruits, green withlightergreen stripes,18–36mm long, 3.0–3.5mmwide,style 7.0–7.5 mmlong,stigma4.5–4.8 unknown, widthunknown;ovary elliptic-oblong,10–10.5mm 1.6–1.8 mmlong;petalsorange, elliptic,apexacute,length trichomes atthroat; sepalsthin,linear, color sameascalyx, 10.0 mmwide,adaxialsurface glabrous orpuberulent with elongated, glabrous, stripedcolor, 17.0–30.0mmlong,0.9– 2–3 pernode,axillary;calyxsalverform,cylindrical-elliptical- long, foldedportion(0.7–)1.0–1.5mmlong.Pistillateflowers appendage round-triangular, glabrous-papillose, 0.1–0.7mm ratio 1.9;anthersfoldedbackwards, 3.4–6.0mmlong,anther date, 4.0–15.0mmlong,3.2–7.0wide,length-to-width 0.8–2.5 mmlong;petalsorange,ovate-rhombic, apexcuspi- at throat; sepalsthin,triangular-linear, colorsameascalyx, 1.0–6.0 mmwide,adaxialsurfaceglabrous withtrichomes salverform, oval,puberulent, solidgreen, 4–9.3mmlong, 2–3 flowersnotmuchlarger thansubsequentflowers;calyx long, with0.27–0.56flowerspermm.Staminateflowers,first diameter, (8–)21–105mmlong;pedicelspuberulent, 1–16mm or rarely lightgreen withdarkersplotches,0.6–1.9mmin long. Inflorescence araceme; pedunclesglabrous, solidgreen attenuate andabaxialbaseacute-truncate, (18-)24–78mm leaflets onpedateleavesbaseoblique,adaxialacute- acute-attenuate, 45–93mmlong,(13–)20–40wide;outer 40–91(–128) mmlong,(12–)21–43wide;sideleafletsbase acuminate-cuspidate; centerleafletbaseacute-attenuate, late andsideleafletslanceolate-elliptic-oblanceolate,apices plex leaveswithcenterleafletelliptic-rhombic-oblanceo- or puberulent, 0.6–1.3mmindiameter, 0–12mmlong;com- depth 44.6%oftotallength;petioluleswingedandglabrous acuminate, 39–98mmlong,40–79wide,averagelobe leaves trilobed,baseauriculate,12–20mmdeep,apex puberulent, 0.5–2.2mmindiameter, 9–45mmlong;simple ther dividedinto2parts),oracombinationofthese;petioles simple, trifoliolate,pedate(sometimeswithouterleafletsfur- often withirregularly-spaced veinsextendingbeyondblade, lent, 3–5mainveins,flush,margins entire-irregular-crisped, surfaces glabrous withveins andmargins sometimespuberu- eter. Leavesimmature membranous,mature coriaceous,both diameter; tendrilsglabrous, solidgreen, 0.3–1.0mmindiam- Anguriacookiana Stems glabrous orpuberulent, solidgreen, 0.9–2.0mmin Plantes deL’Amerique 85 triphyllus, fructovariegato var. 1890, 1902.—TYPE: BAHAMAS. Andros: ConchSound,8May B destroyed fide Jeffrey (1978) ). pedata 242–246 1876.—TYPE: N.J.Jacquin,Sel.Stirp. Amer. Hist.pp. Nov 1899, & Virgin Islands6:267.1925.—TYPE:PUERTO RICO.20 tograph: F!). TYPE: CUBA.Taburete: 16May1839, ottoniana dentata Northrop 556 var. tabula 155 (Schltdl.)C.Jeffrey, KewBull.33(2):352.1978.— Northr., Mem.Torr. Bot.Club.12:69–70t.18. ottoniana Goll 627 Schltdl.,Linnaea24:713.1851. (Schltdl.)Cogn.,Diagn.Cucurb.I:22p.10. Britton Britt &Wils., Sci. Surv. PuertoRico Schltdl.,Linnaea24:758. 1851. (holotype:NY!fide Jeffrey (1978) , pho- . 1763.(holotype!). (holotype:NY!). Sauv., Fl.Cubanap.904. 1873. , C.Plumier, Descriptiondes plate 99 . 1693.(holotype!). Otto 257 Anguria pedata (holotype: Anguria Psiguria

de Anafe, 15Dec 1911, del Río:BañosSanVicente, 12Sep1910, Navas toCampBuenaVista, 650m,23Mar1910, 187 Marianao, Dec1909, 1903, Jul 1900, 1,000 m,08May1986, Tilarán, cloudforest exposedto Atlantic tradewinds,continentaldivide, Harbor, 15Nov1890, 16 Jan1910, asterisk were measured forspeciesdescriptions) hurricane orbymeansofthe Atlantic tradewinds. the Greater Antilles section above),itisnotunreasonable tothinkthatseedsof barium sheetlabel(quotedinthe“ sample. However, becauseofthespecialnotationonher- the foldedanthercharacteristiccouldnotbeconfirmedinthis et al.2010).Thecollectiondidnotcontainmaleflowers,so other herbarium specimenfrom CostaRica,whichgrouped with dominantly intheGreater Antilles, withoneexception, straight anthers( Fig. 6B ). Additionally, Psiguriapedata taxa. variability, thespecieshashistoricallybeensplitintoseveral adjacent positionsalongthestem( Fig. 5A–C ). Becauseofthis change greatly overitslife,andcantakeonvariousformsin est shadeoronslopes,riverbanks,ravines. or dryforests, onrocky, limestone,orclaysoils,indensefor- level to1,200m,insecondary, evergreen ordeciduous,moist winds, ContinentalDivide.”Foundatelevationsfrom sea with labelstating“cloudforest exposedto Atlantic trade and theBahamas( Fig. 4 ), withonecollectionfrom CostaRica flowers orfruits, andtheremainder were sterile. imens examined,70.1%hadmaleflowers,10.4%female part ofthebasemap image. CUBA. Camaguey: LaGloria,30Jan1909, COSTA RICA.Guanacaste:LaChirriparidge, 4kmNEElDosde BAHAMAS. Andros Island:Mangrove Cay, Coppice,LisbonCreek, Examined— Specimens Representative The foldedanther( Fig. 6A ) mosteasilydistinguishes Discussion— Habitat— and Distribution Phenology— Fig. (NY*);Oriente: Antilla, 06Mar1912, Britton &Wilson 70 P. pedata 4 . . Palmer &Riley997 Geographic distributionof Small &Carter8454 samplesinthemolecularphylogeny(Steele from allother Flowering throughout theyear. Ofthe77spec- The leavesofanindividual Camilo 1382 Hitchcock s.n Haber etal.4860 Wilson 11328 P. pedata (NY);Cardenas, 01Sep1903, (NY);Matanzas:mouthofBueyVaca, 28 Aug (F, NY, US);EleutheraIsland:Governor’s were carriedtothecontinentina (NY);IsleofPines:NuevaGerona, 04 . (F). . Throughout the Greater Antilles Greater the Throughout (NY);baseofSierraGuane,26Nov Psiguria (MO*). Psiguria pedata Britton etal.7394 Britton etal.12499 (collections markedwithan Distribution andHabitat Schafer 131 species,whichhave P. pedata Shafer 4418 . Opencircles are Psiguria pedata Britton &Wilson isfoundpre- (NY);Havana: (NY);Sierra (NY);Pinar (NY);Trail ”

Delivered by Publishing Technology to: University of Missouri-Columbia IP: 128.206.9.138 on: Wed, 17 Aug 2011 13:14:52 Copyright (c) American Society for Plant Taxonomists. All rights reserved. tion #2,Z.trifoliolate, variation#3,nearlypedate, AA. trifoliolate,variation #4,nearlypedate,BB.pedate. side leafletmidveinsoff-center, T. trilobed,U.deeply V. shallowlytrilobed,W. deeplytrilobed,X.trifoliolate, trilobed, variation#2,O.deeply variation#3,P. trifoliolate,variation#1,Q.#2,R. trif elliptic, J.adjacentleaves,onesimple, lanceolate,andonebeginningtodivide,K.widetrilobedleaf,L.trilobed, M.deepl E. deeplytrilobed,F. typical trifoliolateleafwithsideleafletmidveinsoff-center, G. trifoliolate withlaterallobesons V–BB: 00 SEL:TXNMCRVSO FPIUI 347 STEELE:TAXONOMIC REVISIONOFPSIGURIA 2010] Fig. P. warscewiczii 5 . . Leaf morphologyvariationinthesixspeciesof . A. adjacent leavesonanimmature branch, B.mature pedate leaf,C.pedatewithsideleafletsfurtherdivided,D.shallowly Psiguria . –: A–C: . P. pedata DE D–E: , P. racemosa FH F–H: , P. ternata ide leaflets,H.rare pedateleaf,I.simple, oliolate, variation#3,S.trifoliolatewith IS I–S: , variation #1, Y) trifoliolate,varia- y trilobed,variation#1,N.deeply P. triphylla TU T–U: , P. umbrosa trilobed, , Delivered by Publishing Technology to: University of Missouri-Columbia IP: 128.206.9.138 on: Wed, 17 Aug 2011 13:14:52 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 4129 Domingo: SantoDomingoCity, near Agua Cave,24Mar1913, Pico DiegodeOcampo,1100 m,17Sep1968, Cabirma delaLoma,700m,28Nov1970, de Torres, 750 m, 30Nov1983, Vallée, Jan1929, Marmelade, 800m,20Dec1925, Botánico, 0–20m,13Jan2007, Banilejo, 800m,01Jun1974, from Juan Aldian, 10Jun1980, Las Mercedes, 400m,22Feb 1969, Banks ofRíoBao,750m,03Jun1968, (NY*); Macoris:Consuelo,15Nov1909, Gastonyetal.161 18 Apr 1964, et al.762 Road, 20Nov1899, Pasto Viejo, 350m,14Oct 1981, San Cristóbal,400m,18Dec1980, Acevedo-Rdgz.&Chinea11659 1996, 14 Dec1941, 11 Jun1981, Romana: RíoCumayasarivervalley, NofBoca de Cumayasa,0–20m, Yuma nexttocave,20–30m,11 Jan2007, above LaPlayita,40–60m,06Feb1981, 05 Jan2007, Pétionville, 350m,15Jun1920, 14 Aug 1903, [Volume 35 1911, appendages. A. folded anthers: the baseofflower, andontherightendsofanthersare theanther SYSTEMATIC BOTANY 348 Yuma, 10–20m,04Feb1981, 18 Sep1985, Nacional LosHaitises,2kmSWdelacasetaMonteBonito,200–300m, El Seibo:LosHaitises,MuelleViejo, 21Jan1980, Clara: Cienfuegos,23May1893, 13383 other speciesof Cordillera Septentrional,300–800m,06May1993, E ofLasSalinas,100–150m,30Oct1985, 400 m,14Mar1980, 2431 Grande, 12 Aug 1920, Britton&Wilson 355 HAITI. GonaveIsland:Étroite, 15Mar1920, PUERTO RICO. Arecibo: Caguana,Río Abajo Forest Preserve, 200–250m, 6. Fig. DOMINICAN REPUBLIC. Azua: LosManantiales,SecciónGalindo, (NY);CiudadTrujillo, 0–25m,07Jan1946, (NY);SouthernBaracoaRegion:30Jul1924, Shafer 10523 (NY);SanDiegode losBaños:05Nov1911, Acevedo-Rdgz. &Axelrod 7781 (US). Two typesofanthersinsidethemalefloraltube.To theleftis Steele etal.1030 Holdridge 889 Mejía etal.1597 Augusto 1469 Mejía &Rimirez 14796 Nash 420 Psiguria (NY*,US);7kmSofLaVega, 16Sep1973, Leonard &11349 (NY, A*); Limestone Hills,Sumidero, 28Jul1912, (NY);Soledad,Jun1941, Mejía 194 Goll etal.627 Loustalot 9459 (F*,NY);Jacmel,0m,Jul1935, . rw b PRS. by Drawn . (NY). (TEX*);Río Abajo Forest Preserve, 340m,18Jan (NY*); LaCiénega,1100–1200 m,13Jul1967, Liogier 21687 (US*);Dept.unknown: Aibonito, Cañónde (NY);Barahona:SierraMartínGarcía, 4km (NY);La Altagracía: 2kmSWofBocade Steele etal.1036 Abbott 4915 Mejía &Zanoni6832 Zanoni etal.10622 Zanoni etal.28069 Liogier etal.32529 Combs 83 (US*);Tijera, J.Diaz,23Nov1899, Psiguria pedata Liogier 14123 (US*);WSabanaHoyos,26Jun 2001, Leonard 8360 Liogier etal.31422 (NY);LaVega: Jarabacoa,1200m, (NY);Santiago: Aug 1940, Liogier 11514 Zanoni etal.10839 Steele etal.1034,1035 (NY);PuertoPlata:LomaIsabel Taylor 166 ; Dept.unknown: Borgne, 0m, (NY);MornesaCabrites,300m, (GH,NY);Sagua,05Sep1903, García etal.578 Liogier 17756 Howard 4792 (TEX*). (NY*);Peravia:9.8kmSW Leonard 3399 Liogier 12694 Smith 10437 (NY);Tortue Island:La Allard 14416 , B.straightanthers:all Leon 11916 (NY);paralleltocoast (NY);Piedra:Firmede Earle s.n (NY*);Piñon,Coamo (NY*);San Cristóbal: (NY*);MataGrande: Bastardo etal.3 (NY*); Pederenales: (NY);Cayey, barrio (NY*);Santiago: Thomas 23 (NY);SaguaLa (NY*); Duarte: (NY);Bocade . (NY*);Santa (NY);Parque (NY). (NY*);Nord: (NY);Santo (NY);Jardín Jiménez 6207 (TEX*);La Rose etal. Clemente (MO); (NY); Shafer Goll

ravines, orclimbingoverrocks andshrubs. in tropical moistordrybroadleaf forests, andonslopes,in Venezuela ( Fig. 7 ). Foundatelevationsfrom 15mto1,750 or fruits, andtheremainder were sterile. examined, 66.7%hadmaleflowers,16.7%femaleflowers been collectedinNovemberandJanuary. Ofthe18specimens other timesoftheyear. A fewfloweringindividualshavealso thickness unknown.Seedsnotseen. exocarp colorunknown,19.0mmlong,12.0wide,wall wide, stylelengthunknown,stigmaunknown.Fruit to-width ratio3.5;ovaryelliptic-oblong,6.5mmlong,3.3 elliptic, apexacuminate,8.8mmlong,2.5wide,length- angular, colorsameascalyx,2.5mmlong;petalsunknown, long, 3.3mmwide,adaxialsurfaceunknown;sepalsthin,tri- salverform elongatedellipse,glabrous, solidgreen, 11.0 mm 0.4–1.0 mmlong.Pistillateflowers3pernode,axillary;calyx 4.4–7.0 mmlong,antherappendagetriangular, papillose, 2.2–7.0 mmwide,length-to-widthratio2.1;anthersstraight, center, elliptic-rhombic, apexacuminate,4.5–13.0mmlong, calyx, 1.5–4.0mmlong;petalspinkwithorangeoryellow dense trichomesatthroat; sepalsthin,linear, colorsameas 5.5(–7.0) mmwide,adaxialsurfaceglabrous withsparseto glabrous orpuberulent, solidgreen, 5.0–9.5mmlong,1.4– than subsequentflowers;calyxsalverformflask-shaped, per mm.Staminateflowers,first2–3flowersnotmuchlarger pubescent, 2.0–13.5mmlong,with0.5–0.74(–0.83)flowers green, 1.1–2.0mmindiameter, 90–225mmlong;pedicels terminate corymb;pedunclesglabrous orpuberulent, solid depth average67.3%oftotallength.Inflorescence aninde- apex acuminate,115–219 mmlong,32–70wide,lobe auriculate-truncate-rounded-acute, base7–30mmdeep, in diameter, (3–)22–58mmlong;simpleleavestrilobed,base blade, simple;petiolesglabrous orpubescent,1.2–3.0mm ular-wavy, withirregularly-spaced veinsextendingbeyond times puberulent, 3–5mainveins,flush,margins entire-irreg- Leaves membranous,bothsurfacesglabrous withveinssome- diameter; tendrilsglabrous, solidgreen, 0.5–1.6mmin diameter. 2. part ofthebasemap image. morphology of Discussion— Habitat— and Distribution Phenology— Stems glabrous orpuberulent, solidgreen, 1.3–4.7mmin Fig. Psiguria racemosa 10130 1978.—TYPE: VENEZUELA. Aragua: Guamitas, 7 . . Geographic distributionof (holotype:VENfide Jeffrey (1978) ). Flowering April through July, andperhaps Compared toother speciesof P. racemosa C.Jeffrey, KewBulletin33(2):347. isfairlyconsistent,trilobed and otnna Clmi and Colombia Continental Psiguria racemosa Psiguria . Opencircles are , theleaf Williams s , Delivered by Publishing Technology to: University of Missouri-Columbia IP: 128.206.9.138 on: Wed, 17 Aug 2011 13:14:52 Copyright (c) American Society for Plant Taxonomists. All rights reserved. and LaRinconada,550m,13Nov1979, 06 May1983, wood, 1,000–1,200m,17May1918, Cabello, 0–30m,02Jul1913, 25 Jan1961, 260 m,Jul1978, 14999 Selvas deRanchoGrande,Parque Nacional,1,100m,17May1942, 1937, Huarapiche, 30 Apr 1979, 11422 Zulia: Bolívar, 6kmEofElPensado,150–200m,27 Apr 1982, 3 3. 1984, 109898 Palavecino, betweenSarare &El Altar, 350m,02Jun1974, Catuche, NofCaracas,1,200–1,800m,01Jun1913, Barriga 13400(NY*). Cundinamarca: Sasaima,QuebradaLaMaría,1,750m,07Jan1950,García- asterisk were measured forspeciesdescriptions) 349 ( cally withayellowbase),maleflowerpedicelaveragelength petal color( acteristics thatdistinguishthesetwospeciesincludemale morphs of with STEELE:TAXONOMIC REVISIONOFPSIGURIA speckles in peduncles andcalyxtubesthatare lightgreen with darker as opposedtotheorangeororange-red flowerpetals,and petals, andpedunclescalyxtubesthatare solidgreen, P. racemosa have, simple,trilobedleaves.Thecharactersthatseparate and The geographicrangeof membranous, butwithvariabledepthoflobes( Fig. 5D,E ). 2010] margins entire-irregular-wavy, with infrequent irregularly- brous, veins flushorseldomabaxialmain veinsprominent, 0.8–1.7 mmindiameter. Leavescoriaceous,bothsurfacesgla- ear splotches,2.0–5.6mmlong; tendrilsglabrous, solid green, Anguriagloriosa Anguriagrandiflora flowers alongthepeduncle( Anguriakunthiana did notstrongly supportthedistinctionof obvious morphologicaldifferences, themolecularphylogeny mm; two speciesmaybecomebetterdefined. more collectionsandmolecular study, thelinesbetweenthese and onlyfiveincludedinthemolecularphylogeny).With als of P. triphylla P. racemosa The geographicrangeof Stems glabrous, solidgreen orlightgreen withdarker, lin- Psiguria ter VENEZUELA. Antioquia: 22Jun1946, COLOMBIA. Bolívar:nearSincé,20 Apr 1963,Romero 9685(MO*); Examined— Specimens Representative photograph: F!). 1892, IV p.366t.25.1895.—TYPE: Localityunknown.1891– (holotype: W; photograph:F!). TYPE: BRAZIL.Maynas: Yurimaguas, 1831, (holotype fide Jeffrey (1978) ). 2: 26.1846.—TYPE:t.2inVell., Fl.Flumin. p.10.1835. 33(2): 354.1978. BRAZIL. Dec,1836, (NY*);Dept. unknown:5kmWofJusepin,MonagasontheRío P. warscewiczii (US*);Bolívar:Piar, LaCamilera,40kmWofElManteco,250– Tamayo 432 Aymard &Ortega2527 P. triphylla P. triphylla (NY);Portuguesa: Araure, 5kmNEof Agua Blanca,250m,05May P. racemosa Moore 659 (Steeleetal.2010).There havebeenfewindividu- from theothertwoare itsdistinctivepinkflower Steyermark 88837 P. umbrosa P. triphylla =5.6mm; P. racemosa Stergios 5572 (US). Delascio &Liesner7096 ; however, inthisarea, thesimple,trilobedleaf = S.Moore, Trans. Linn.Soc. Lond. ser. 2.Bot. nata collected(only18examinedinthisstudy, > Schltdl.,Linnaea24:762.1851.—TYPE: Cogn.Diagn.Cucurb.fasc.Ip.11. 1876.— inVenezuela, andallthree have,ormay 0.75flowerspermm).Despitethesefairly (holotype:NY!). n and (M.Roem.) C. Jeffrey, KewBulletin havenotbeenseen. Additional char- Anguria ternata Seigler 11153 =pink; P. triphylla (NY);Guanare, terrenos delaUNELLEZ, (NY);Tachira: between Aguas Calientes Humbolt 8978 Pittier 6426 P. racemosa (NY*);Carabobo:ElPalito,nearPuerto P. warscewiczii Psiguria racemosa P. racemosa Pittier 7854 P. triphylla (MO);Faldassurdel Avila, Aug (US*);EofMiamo,400–550m, =absent),anddensityof (US*);Caracas:LowerCatuche Steyermark etal.120279 overlapswith (collections markedwithan (holotype:Bdestroyed; M. Roem.,Syn.Mon. Haught 4898 (US);Federal:bosquede = . Pittier 6294 =deepred, typi- < 0.75flowersper P. racemosa alsooverlaps Steyermark etal. (S; Aragua: (US); Poeppig 2416 (NY*);Lara: P. umbrosa (MO*); Bunting from Pittier

guay ( Fig. 8 ). Foundatelevationsfrom 170mto1,800 or fruits, andtheremainder were sterile. examined, 71.2%hadmaleflowers,19.2%femaleflowers ing peakfrom Octoberthrough February. Of the73specimens mm long,4.5–4.9wide,1.9–3.0thick. 10–18 mmwide,wall0.7–2.0thick.Seeds44–54,6.5–7.9 green, sometimesspeckled withlightergreen, 22–44mmlong, style 3.3–5.0mmlong,stigma5.0–7.0(–12.0)long.Fruit ovary elliptic-oblong,12.0–20.0mmlong,4.2–7.0wide, 8.0–16.0 mmlong,3.5–9.0wide,length-to-widthratio2.1; long; petalspink,obovate,apexrounded orrarely cuspidate, throat; sepalsthin,linear, colorsameascalyx,2.3–3.7mm adaxial surfaceglabrous orpuberulent withtrichomesat glabrous, solidgreen, 19.0–32.0mmlong,2.5–5.2wide, with reduced leaves;calyx salverform,cylindrical-elliptical, flowers 2–4pernode,axillary, butattheendsofbranches triangular, glabrous-papillose, 0.2–1.0mmlong.Pistillate anthers straight,4.4–8.0mmlong,antherappendageround- ers 40.0,25.0,and11.0 mmwide),length-to-widthratio1.2; and 22.0mmlong),2.4–7.2wide(averagefirst3flow- rounded, 3.0–10.5mmlong (averagefirst3flowers62.0,48.0, rose, salmon-rose, ordeep-rose, orbicular-obovate, apex thin, linear, colorsameascalyx,1.0–8.5mmlong;petalspink, wide, adaxialsurfaceglabrous withtrichomesatthroat; sepals green withdarkerspeckles,5.0–13.0mmlong,2.0–6.4 drical-elliptical, glabrous orpuberulent, solidgreen orlight much larger thansubsequentflowers;calyxsalverform,cylin- 0.25–1.00 flowerspermm.Staminateflowers,first2–3 long; pedicelspubescent,(0.1–)0.5–1.6(–4.0)mmlong,with darker splotches,1.5–3.0mmindiameter, (78–)155–293mm cles glabrous orpuberulent, solidgreen orlightgreen with long, 29–97mmwide.Inflorescence acompactraceme;pedun- acute-attenuate andabaxialbaseacute-truncate, 80–172mm long, 45–115 mmwide;sideleafletsbaseoblique,adaxial acuminate; centerleafletbaseacute-attenuate,95–200mm tic, oblique,andsometimeswithsmall,laterallobe,apices rhombic-oblanceolate-spatulate andsideleafletsovate-ellip- in diameter, 2–35mmlong;complexleaveswithcenterleaflet 14–80 mmlong;petiolulesglabrous orpuberulent, 1.2–2.0mm bifid orpedate);petiolesglabrous, 1.3–3.3mmindiameter, spaced veinsextendingbeyondblade;trifoliolate(orrarely part ofthebasemap image. Fig. 8. 8. Fig. Dsrbto ad Habitat— and Distribution Phenology— Geographic distribution of Flowering throughout theyearwithflower- Bolivia, Brazil, Perú, andPara- Psiguria ternata . Opencircles are s , Delivered by Publishing Technology to: University of Missouri-Columbia IP: 128.206.9.138 on: Wed, 17 Aug 2011 13:14:52 Copyright (c) American Society for Plant Taxonomists. All rights reserved. angular) and with three subsequentflowersonthesameinflorescence (small). P. triphylla cate great resources tomakingagiantdisplay. where tofindtheflowers, theplantnolongerneedstoallo- plants (L.E.Gilbert,pers.comm.).Oncethepollinatorknows linating butterfliestothelocationsofthesewidelyseparated [Volume 35 esized tobeanadaptationforattractingandtrainingthepol- than subsequentflowers( Fig. 9 ). Thischaracteristicishypoth- on theinflorescence, which canbesixtoseventimeslarger ternata climbing overothervegetation,andonsandyorclaysoils. dense shadeorlightgaps,onroadsides, riverbanks,orslopes, blands. Maybefoundindisturbedorsecondaryforests, in or drybroadleaf forests, floodedgrasslands,andxericshru- in tropical evergreen, deciduous,orsemideciduous,moist SYSTEMATIC BOTANY 350 P. ternata pedate), coriaceousleavesandpinkflowerseasilydistinguish P. ternata membranous leavesandorangeflowers.Distinguishing Also, maleinflorescences of leaflets are typicallycentered, thoughnotalways( Fig. 5P–S ). off-center ( Fig. 5F–H ), whereas midveinsin more oftenin have predominantly trifoliolate, coriaceousleaves,although spaced flowersandpinkpetals.Maleinflorescences of flower clusterhasatriangularshape( Fig. 10A ) withloosely Fig. The geographicdistributionof Discussion— Fig. 10 9 isthelarge sizeofthefirsttwoorthree maleflowers . . . . rm from rm from Comparison ofthefirsttwoflowers andafew Comparison oftheflowerclusters on P. triphylla P. ternata P. triphylla The mostdistinctivecharacteristicof P. umbrosa B hrzna ellipse). horizontal (B. P. umbrosa , themidveinsof sideleafletsare isnotquiteasstraightforward. Both , whichhassimple3-to5-lobed, P. ternata . Consistenttrifoliolate(rarely Psiguria ternata are racemose,andthe Psiguria ternata Psiguria ternata P. triphylla overlapswith Psiguria (A.tri- (large) side Nacional Amboró, 500m,21Jan1988, (NY); Florida,Bermejo,700m,08Dec1988, NE ofjunctionwithRíoBermejo,800m,16Feb1988, 7 kmSEofNaranjillos,480m,30Sep1990, Puerto PailasHwy, 320m,07Jan1998, (NY); SantaCruz: Andrés Ibáñez,2.8kmSEofturnoff from SantaCruz to 1.5 kmSWofMontero, 295m,12Jan1987, Jan 1983, Gerais: Capinópolis,Chácara,15Dec1995, 30 kmSEofGoiásVelho, 700m,21Jan1966, NE ofCatalão,875m,23Jan1970, Jussari, SerradoTeimoso, 300m,31Jan1999, retera enoestedesendero, 618m,10Jan2008, 2008, derecho ladodelsendero queestásurdepuenteala Mistad,317m,09Jan 1995, Dec 1995, 1988, 30 Oct1984, et al.1762 (NY); Nor Yungas, km12Coroico toCaranavi,1,015m,23Jan1983, de Villa Barrientos19kmhacia Arapata, 1125 m,04Jan1990, Yolosa, 1,150m,30Oct1984, 05 Oct1984, 1884 14 Dec1988, 1988, asterisk were measured forspeciesdescriptions) Curitiba, SeteQuedas,Guaira,12Dec1977, Serra doCachímbo,425m,12Dec1956, 900 m,Dec1929, (F, MO); SanRoque,Feb1930, Tarapoto, banksofChumbazariver, 830m,25Feb1947, Chazuta, 300–400m,18May1986, Río Huallaga,06Oct1970, 9629 Parque NacionalManu,CochaCashuStation,350m,13Jul 1984, (F*); 39kmSWofPuertoMaldonado, 28Jan1989, Manu, Parque NacionalManu,400m,26Oct1986, Requena, 170m,04Dec1980, 220 m,16Dec1964, 1986, al.1488 Grosso, RibeirãodaOn Mato Grosso doSul,16Mar1985, 19 Jan1983, Ceará: Crato,Chapadado Araripe, Parque Nacionaldo Araripe, 820m, Parque SeteQuedas,10Jan 1979, 15 Sep1982, 1986, 1387 7808 Ariquemes, 11 Oct1979, & Wilson 4245 et al.7405 Velho, reservatório daUsinaHidroelétrica deSamuel,08Jun1986, or subspeciesinthefuture. ular studywillsupportasplitofthesecladesintotwospecies split intotwotaxa.Perhaps additionalcollectionsandmolec- no morphologicalcharacteristicswere identifiedtosupporta the tropical, dryforests ofsouth-centralBolivia. However, and northernBolivia,theotherincludedsamplesfrom three samplesfrom thetropical, moistforests ofPerú, Brazil, two subcladeswere strongly supported.Oneclade included Psiguriaternata petals thatare deepred, typicallywithayellowbase. horizontal ellipse( Fig. 10B ) withtightlyclustered flowersand P. triphylla The molecularphylogenystrongly supportedthecladeof BRAZIL. Amazonas: Camatian,24Jan1949, BOLIVIA. ElBeni:Ballivián,SofMisiónFatima,300–500m,24May Examined— Specimens Representative PERÚ. Cusco:Calca,Quebrada,1,031m,26Jan2003, PARAGUAY. SanPedro: 27 Apr 1958, (NY);LaPaz:Nor Yungas, 21kmNWofChuspipata,1,800m, (F*);SanMartin:MariscalCáceres, Tananta, margen izquierda del (MO); RíoTambopata, 2kmofColpadeGuacamayos,280m,10Nov (NY*). Seidel &Schulte2303 Steele etal.1040 Beck etal.16469 Guillen etal.3178 Mendon Núñez 6511 (NY);DF:FazendaMariaPereira, RegiãodoMesquita,GO,05Feb (F, MO);Nor Yungas, onroad from Coroico toCaranavi,1,150m, Fernández 8054 (NY);FortePrincipedebeira,localBaia,05Jan1962, Nee 46377 are spicate,andtheflowercluster isshapedlikea Nee 37179 Plowman 12738 Nee &Solomon30243 Foster 8808 Solomon &Escobar12460 ç (NY*);Minera a &Alvarenga 608 (MO);Huánco:Pachitea,NofPuertoInca,250–300m, Williams 6514 samples(Steeleetal.2010).Within thisclade, (NY);Velasco, CampamentoRefugio,180m,11 Feb Schunke 6675 (TEX*);RíoPirai,puenteTaruma; 1kmsurdelacar- (F);Cercado, Casarabe,51kmEofTrinidad, 200m, (NY*);Rurrenabaque, 1,000ft,28Nov1921, (NY);300m,16May1991, (MO);Loreto: Contamana,QuebradadeMaquia, (NY);Reyes:SanPedro, 25Oct1921, Zarucchi etal.2678 ç (NY);Pando:NicolásSuárez, cerca deCobija,07 inha, Poconé,Cuiabá,12Feb1980, Schunke 4482 (F*,NY);Curitiba: Antonio João,Campestre, Nee &Solomon30243 Vásquez &Jaramillo872 ç Williams 6946 ão Taboca atmassangana,35kmWSWof (F); Ucayali:Purús, RíoCuranja,cerca la (US);Goiás:ContraforteCentral,20km (F, TEX,US);Sud Yungas, 450m,01Jan (MO);RíoUcayali,1horasurcando de Knapp &Alcorn7315 Irwin etal.25200 Bernardi 19433 Hatschbach &Zelma49096 (MO,NY);Nor Yungas, 9kmNof (NY);SanMartin,hillstoNWof Nee 47832 Saldias 134 Woolston 960 Nee 38959 Nee 33434 Pires etal.6158 (NY). Nee 37059 (US); Alto RíoHuallaga,360– Macedo 4071 (collections markedwithan Irwin etal.11903 Steele etal.1043 Thomas etal.11882 Hatschbach 40595 Fróes 23975 (MO,NY);Nor Yungas, (NY);Rondônia:Porto (Y) Adé Ibáñez, Andrés (NY*); Foster &d’Achille12050 Smith etal.1644 (US*);SerraDourada, Gentry etal.73937 (MO); Madre deDios: (TEX*);Florida,1km (NY);Sara,260m,20 (NY);Santiesteban, (NY);Ichilo,Parque Nee &Saldias36321 (NY*,US); (MO);SanMartin, Woytkowsiki 35180 (MO*,US);Pará: Valenzuela etal. (Y; Paraná: (NY); (NY);Bahia: (NY);Minas Beck 17361A (US);Mato (TEX*). White 1541 Macedo et Rodrigues Cardenas (NY*); Ferreira (NY*); Hassler (NY); Foster Besse (F);

Delivered by Publishing Technology to: University of Missouri-Columbia IP: 128.206.9.138 on: Wed, 17 Aug 2011 13:14:52 Copyright (c) American Society for Plant Taxonomists. All rights reserved. Anguriadiversifolia Anguriatriphylla Anguriaboissieriana Anguriapachyphylla Anguriatreslingiana Anguriaoblongifolia Anguriaschomburgkiana Anguriavogliana Angurialongeracemosa Anguria dunlapii 351 STEELE:TAXONOMIC REVISIONOFPSIGURIA 4. 1061 comunidad nativadeColumbiana,250m,22Feb2000, 2010] Anguriatabascensis Angurialongipedunculata Anguriapallida Anguriaelliptica Psiguria triphylla (NY). TYPE: PERU. Alta Verapaz, Nov1888, Casas, Fontqueria52:4.1998.—TYPE:GUATEMALA. 16(1): 10.1891. 6149 p. 670.1881.—TYPE:ECUADOR,Chimborazo graphs: F!,MO!). (1978)). Wunderlin fide Jeffrey (1978) ). 236. 1912.—TYPE:COSTA RICA. online photographU!). TYPE: SURINAME.Jul1908. 1510 GUATEMALA. Livingston:RíoDulce,Mar1889, GUYANA. (1978) ; photograph:F!). TYPE: SURINAME. 347. 1978. Jeffrey (1978) ; photographs:F!,MO!). VENEZUELA. Maracay, 1928, Wunderlin (1978) ; isotype:onlinephotograph US!). margin offorest, Apr 1914, 1929.—TYPE: PANAMA. Yaviza, southernDarien,on US!; photographs:F!,MO!). Changuinola Valley, 3Mar1924, 339. 1992.—TYPE:PANAMA. BocasdelToro: FarmSix, 1929. Catemaco, Apr 1845, Phytologia 38(3):219.1978.—TYPE:MEXICO.Veracruz: US fide Wunderlin (1978) ; onlinephotograph!). type: NY!;isotype:Kfide Jeffrey (1978) ). TYPE: MEXICO.Tabasco, 4Jul1889, Jeffrey (1978) ; photographs:F!,MO!). 21. 1876. 6205 1881—TYPE: ECUADOR.Chimborazo.Jun1860, triphylla type: US!;isotype:NY!). TYPE: TRINIDAD. Arima, 4Mar1921, (holotype:K). (holotype:onlinephotographUS!). (holotype:Kfide Jeffrey (1978) ; photograph:F!). Psiguria dunlapii var. Anguria triphylla Psiguria longipedunculata Cogn.,Diagn.Cucurb.I:22.1876. Schomburgk 1254 Britton,Bull.Torr. Bot. Cl.50:54.1923.— Standl.,FieldMus.Publ.Bot.Ser. 4: 298. Suess.,FeddeRep.30:278.1932.—TYPE: var. Cogn.,Donn.Sm.Bot.Gaz.(Crawfordsville) Psiguria diversifolia Donn.Sm.,Bot.Gaz.54:236–237.1912.— pallida Pulle,Rec.Trav. Bot.Néerl.6:289.1909.— Cogn.DC.Mon.Phan.IIIp.673.1881.— Donn.Sm.,Bot.Gaz.(Crawfordsville) 54: Cogn.,Bot.Gaz.16:9.1891.—TYPE: Ruiz 34096 Pittier, J.Wash. Acad. Sci.19:185. acuminata Schltdl.,Linnaea24:765.1851.—TYPE: (Miq.)C.Jeffrey, KewBulletin33(2): Kappler 1728 Cogn.,DC.Mon.Phan.III.670. Cogn.Diagn.Cucurb.fasc.Ip. (Standl.)R.J.Hampshire, Novon2: Galeotti s.n Turckheim 1414 Miq.Linnaea19:136.1845.— Pittier 6582 (holotype:G-BOIS;photo- Cogn.,DC.Mon.Phan.III (holotype:Bdestroyed fide Vogl 302 (Cogn.)C.Nelson,Fern.& (holotype:Ufide Jeffrey Tresling 252 Tonduz 11535 Dunlap 490 (Cogn.)Wunderlin, . (lectotype:BRfide Rovirosa 519 (holotype:USfide , (holotype: Mfide Britton 2084 (holotype:US! Graham &Schunke (holotype: (holotype: (holotype: Psiguria (holo- (holo- Spruce Spruce Smith used todistinguish widely distributed geographically( Fig. 11 ). Forcharacters sizes toanassortmentoftrifoliolate profiles. Itisalsothe most range from simpleandelliptictovarioustrilobedshapes and most variableleafmorphology ( Fig. 5I-S ). Mature leaves tion, andonlimestone,sandy, orclaysoils. roadsides, riverbanks,orslopes,climbingoverothervegeta- ary forests. Maybefoundindenseshadeorlightgaps,on the uppercanopiesorclearingsofbothprimaryandsecond- moist ordryforests, cloudforests, andfloodedgrasslands,in ical andsubtropical evergreen, deciduous,orsemideciduous, ( Fig. 11 ). Foundatelevationsfrom sealevelto2,100mintrop- range of female flowersorfruits, andtheremainder were sterile. specimens examined,75.0%hadmaleflowers,9.1% long, 4.2–5.9mmwide,1.8–3.8thick. mm wide,wall0.7–3.0thick.Seeds32–46,7.7–10.0 stripes, someturningyellowwithage,30–45mmlong,15–24 stigma 6.1–10.2mmlong.Fruits, green withlightergreen 14.0–22.0 mmlong,3.0–5.8wide,style4.6–9.0 9.5 mmwide,length-to-widthratio1.6;ovaryelliptic-oblong, rhombic-oblanceolate, apex cuspidate,4.5–9.5mmlong,2.5– 1.0–2.3 mmlong;petalsred, orange-red, orscarlet,orbicular- linear, colortypicallysameascalyx,butsometimesdarker, glabrous withtrichomes atthroat; sepalsthin,triangular- green, 11.0–41.0 mmlong,1.4–6.5wide, adaxialsurface node, terminal;calyxsalverform,cylindrical,glabrous, solid brous-papillose, 0.1–1.2mm long.Pistillateflowers2–5per 3.4–8.5 mmlong,antherappendageround-triangular, gla- 2.0–10.0 mmwide,length-to-widthratio1.6;anthersstraight, lar-rhombic-flask-shaped, apexcuspidate,2.0–12.0mmlong, with yellowcenter, orange-red, strong red, orscarlet,orbicu- calyx, butsometimesdarker, 0.6–2.7(–4.8)mmlong;petalsred throat; sepalsthin,triangular-linear, colortypicallysameas adaxial surfaceglabrous with sparsetodensetrichomesat puberulent, solidgreen, 5.5–15.0mm long,1.0–4.8mmwide, often withathickenedbase,rarely cylindrical,glabrous or larger thansubsequentflowers; calyxsalverformcylindrical, ers permm.Staminateflowers,first2–3flowersnotmuch absent orlengthlessthan0.3mm,withgreater than0.75flow- (0.6–)1.1–4.6(–8.0) mmindiameter, 63–395mmlong;pedicels indeterminate corymb;pedunclesglabrous, solidgreen, mm long,18–124wide.Inflorescence anapedicellate, wide; sideleafletsbasecuneate-acute,rarely oblique,55–225 leaflet baseacute-attenuate,71–305mmlong,22–132(–177) lets ovate-elliptic-obovate,apicesacuminate-cuspidate;center with centerleafletelliptic-obovate-oblanceolateandsideleaf- brous, 0.9–3.6mmindiameter, 2–48mmlong;complexleaves wide, lobedepthaverage53.2%oftotallength;petiolulesgla- deep, apexacuminate,75–184mmlong,(22–)40–76(–178) or trilobed,baseauriculate-rounded-acute, base5–30mm in diameter, 8–75mmlong;simpleleaveslanceolate,elliptic, olate, oracombinationofthese;petiolesglabrous, 1.0–5.3mm ularly-spaced veinsextendingbeyondblade,simple,trifoli- margins entire-irregular-crisped-revolute, seldomwithirreg- flush orseldomabaxialandadaxialmainveinsprominent, diameter. Leavescoriaceous,bothsurfacesglabrous, veins in diameter;tendrilsglabrous, solidgreen, 0.3–3.1 mmin older, woodystems,solidgreen orbrown, 1.1–7.0(–32.0)mm Stems puberulent atnodes,otherwiseglabrous, flakingon Discussion— Habitat— and Distribution Phenology— Psiguria Flowering throughout theyear. Ofthe297 f h the Of , from southernMexicotonorthernBolivia P. triphylla Psiguria from theother species,seethe species, hogot h Continental the Throughout P. triphylla a the has Delivered by Publishing Technology to: University of Missouri-Columbia IP: 128.206.9.138 on: Wed, 17 Aug 2011 13:14:52 Copyright (c) American Society for Plant Taxonomists. All rights reserved. part ofthebasemapimage. taining approximately 383flowerscars. undergoing molecularandmorphologicalstudymayleadto than rotate (L.E.Gilbertpers.comm.). Additional collections the femalecorollas are typicallyreflexed downward rather leaf morphs. Additionally, although rarely seeninthefield, rounded ( Fig. 5K ) thanseenintheothertrilobed but thelobingismuchmore shallow, andlobetipsare more The leavesofthesespecimensare simple,trilobed,andwide, clade, itdoesnotexplaintheratherlongbranchleadingtoit. another, artificiallyincreasing thebootstrapsupportfor ples includedinthemolecularphylogenyare clonesofone Catemaco, Veracruz, Mexico. Although fourofthesixsam- One exceptionmaybethespecimensfrom LosTuxtlas and morphology donothelptoexplainthesemoleculargroups. cal variationinmaleflowers,andthegreat variationinleaf samples andsomelongbranches.Thelackofmorphologi- supported clade.Within thiscladewere several groupings of of the28 had beenfloweringfor7–13moatthetimeitwascollected. that thisinflorescence (with approximately 383flowerscars) cle. Typically onlyoneortwoflowersopenperdayimplying [Volume 35 P. triphylla (as manyasthree permillimeter). Figure 12 showsatypical pedicels, andahighdensityofflowersonaninflorescence Psiguriatriphylla discussions underthosetaxonomicdescriptionsorthekeys. SYSTEMATIC BOTANY 352 Fig. Fig. In themolecularphylogeny(Steeleetal.2010),allbutone 12 11 . . . . Psiguria triphylla inflorescence withflowerscarscoveringthepedun- Geographic distributionof Ifoecne f of Inflorescence ismosteasilyidentifiedbyflowersthatlack Psiguria triphylla samplesformedamoderatelywell- Psiguria triphylla , nearly14cmlong,andcon- . Opencircles are P. triphylla

(LL); Tikal NationalPark,04Mar1961, 3363 39121 Steyermark48982 Huehuetenango: SierradelosCuchumatanes,1,500–1,600m,17Jul1942, (NY*); Northwest:Waini River, 06Feb1922, El Porvenir, 1,300m,06Mar1949, Region of Ajaxá, 330m,23Feb1941, Station, 200–300m,24Oct1998, 12 Jun1995, 25 kmNEofDanlí, 16May1982, Montaña LaCumbre, 600m,21Mar1962, (GH); Colón:3km EofTrujillo, 09Jun1980, Boom&Gopaul7305 (US); Dept.unknown:UpperMazaruni Riverregion, 500m,15 Apr 1987, Demerara-Berbice: 83miBartica-Potaro Road,400ft,24Jun1933, Türckheim 775 19041 Hahn3652 Grubbetal.1276 15 Nov1998, (MO); LaPaz:Larecaja, 1,200m,23Jan1988, Jan 1946, Toledo: CohuneRidge,15Mar1945, asterisk were measured forspeciesdescriptions) samples. the designationofavarietyorsubspeciescontainingthese Aug 1989, 1973, Mathias&Taylor 5232 1994, Cordillera deTilaran, 1,500m,01 Aug 1996, the small-stemmedformisfound. tion isinnortheasternVenezuela andinTrinidad where only throughout thegeographicrange.However, anotableexcep- mm). Collectionsofwidelyvaryingsizeshavebeenmade leaves, whereas othershaverobust, sturdy stems(upto32 individuals havesmallstems(1.0mm)andalmostdelicate in stemsizeandoverallrobustness. Somemature, flowering 1974, Lawrance707 (NY*). 4529A 01 Dec1995, Yungas, 31Oct1984, 20–300 m,12Feb1988, Nariño: Espisella,Fumaco,20Jun1951, Meta: Mesetas,517–600m,29Feb1988, 207 Barriga 11729 (MO); Cundinamarca: WofGuaduas,1,040–1,320m,03Nov1945, Croat 22130 Robles1728 1986, 18 Feb1997, Puerto Viejo, 100m,27Mar1983, 1985, Campos144 Steele&Fernández1008 Utley&2804 1985, 11 Aug 1935, Estación BiologicaLasCruces, 1,000–1,100m,08Mar1996, Psiguriatriphylla (US);ElValle: RíoDiguaValley, 825m,02 Apr 1939, GUYANA. Morawhanna:BarimaRiver, 14 Jan1920, GUATEMALA. Alta Verapaz: Cubilquitz,350m, Apr 1901, FRENCH GUIANA.Saül:MontsLaFumée,200–300 m,26Jul1987, HONDURAS. Atlantida: Peru, 01May1970, BOLIVIA. Beni:Ballivián,1025m,25Sep1984, BELIZE. Cayo:Millionario,1,800ft,30May1973, Examined— Specimens Representative ECUADOR. Esmeraldas:BilsaBiologicalReserve,400–600m,15Nov COLOMBIA. Boyaca:I30M-NofBogotá,3,000ft,23Mar1933, BRAZIL. Acre: Brasiléia,07Nov1991, COSTA RICA. Alajuela: NofSanRamón,550–1,150m,08 Aug 1975, (NY*);Petén:DosLagunasinZapotal,26Oct1960, Stevens 23940 Haber &Bello2039 (F);NWofLakeIzabal,500–600m,11 May1966, (NY*);Bélizon,200–400m,03 Aug 1993, Monsalve 812 Holm-Nielsen etal.6964 Gentry etal.9134 Pitman &Clark937 (US*);Serra dosCarajás,450m,17Jun1982, Gentle 5469 Cerón &Factos7619 (NY);trailtoMt.Galbao,250–300m,18 Aug 1988, (MO);Puntarenas: BuricaPeninsula,50–200m,22Feb1973, (MO*);Monteverde, 1,500m,24Dec1985, (MO);Guanacaste: LaCruz de Abangares, 1,400m,15Jul (A);Caqueta:10kmSofSanJosedeFragua,320m,11 Jan Haught 1874 (US);ElMeta:Villavicencio, Jun1930, Gonzales etal.1756 Aguinda etal.250 Oliveira etal.705 (US);Chirriacté,900m,09 Apr 1941, Acevedo-Rdgz. &Cedeño7289 (NY). (F);Izabal:RíoJuyamá,50–150 m,08 Apr 1940, (MO). (MO);Parque NacionalTortuguero, 40m,14Mar1988, (F*);HotelVilla Blanca, 1,100–1,200m,25Jul2005, (NY*). (LL*);MangaCamp,30 Apr 1948, Solomon &Nee12650 Cárdenas 1247 (F);Napo:road Papallacta-Baeza,2050m,06Jun alsoincludesindividualsthatare different (MO);Choco:Pizarro, 16Nov1985, (MO);Heredia: LaSelva,OTSFieldStationonRío (TEX*);Cartago:Turrialba, 1,140m,16Jul1993, (F*);Valle delCauca:BajoCalima,100m,22Mar (MO);Guayas:Santiago-Zamora,22Jul1960, (MO*);JatúnSachaBiologicalStation,450m, (NY*);Cantón Archidona, 800–1,000m,23 (F);Pichincha:Borja,5,600ft,12 Aug 1960, (NY);Pará:Belém,17Jun1944, (MO);Cerro Coronel, 10–100m,24Jan Chacon 601 Burnham 1761 Steyermark 37069 Sánchez 184 (MO);Santander:Caquetá,100–700m, Standley 88231 Gentle 5272 Romero 2794 Callejas &Marulanda6033 Lundell 16872 Ferreira &Ming145 (NY). (F);Limón:Talamanca, 500m, (collections markedwithan Molina 10545a (US*);Tiputini Biodiversity Krings 59 de laCruz1297 Saunders 309 Solomon 17679 Mori etal.23084 (MO);Olancho: Montaña (F);Río Aguarico, 235m, Barkley &Hernández40227 , (LL*);San Antonio, 03 (F);SanMarcos: Finca (F). Schmitt &141 (F);Ríosucio:Urabá, (F*). Herb. Nac.Colombia Standley 91695 Sperling etal.6217 (LL*);Retalhuleu: Gentry 7692 Jones &Facey3361, Haber 3887 Gentle 6520 Killip 34737 (F);El Paraíso: Hitchcock 17499 (TEX);Cortés: Contreras 1561 Krings 246 (NY*);Upper (NY*);Feijó, (MO*);Nor Espina 1833 (NY). Steyermark Mori etal. Tutin 255 Baldwin (MO*); , (LL). , Garcia- (MO); (MO); (F); (F); (F); von

Delivered by Publishing Technology to: University of Missouri-Columbia IP: 128.206.9.138 on: Wed, 17 Aug 2011 13:14:52 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 19 Dec1978, 300–400 m,13Feb1967, 1,500–1,600 m,23Jun1976, Río Monzón,672m,06Mar1962, Cenepa, Huampami,200–250m,1978, 18738 (MO). (MO*); Veraguas: beyondTres BrazosRiver, 650m,24Jul1974, Colorado Island,CanalZone,WheelerTrail, 17Sep1970, NW ofCerro Jefe,650–750m,08Nov1979, Campana, 120mdelMotelSulin,3,065ft,26Dec1977, El Llano-CartiRoad,350m,15Feb1975, (NY); 1–2mibeyondGoofyLake,01Jan1972, Lewisetal.2217 Judziewicz4520 McPherson7085 00 SEL:TXNMCRVSO FPIUI 353 STEELE:TAXONOMIC REVISIONOFPSIGURIA National Park,1,400m,06Jun1992, Los Zapatos,1,200–1,500m,25 Apr 1982, 2010] Liesner23801 1961, 1982, 12145 Steyermark61005 Brittonetal.418 Cowan&Forster1395 Britton 2084 250–350 m,10Mar1956, Texas greenhouse, 15Feb2008, 16951 2008, 18 Jan2008, Pucuraquillo, 08Mar1981, Río Amazonas SofIquitos,18 Aug 1972, (MO*); Previsto, Yurac Canyon,420m,09Oct1962, Jun 1981, Davidseetal.20329 550 m,02 Apr 1983, (MO); Ocosingo,170m,14Jun1985, Bárbara: SanPedro Sula,Jul1887, (F); Sipaliwini:Frederik Top, 500m,20 Aug 1963, (NY); Rheno-Trai.: LelyMts.,550–710m,26Sep1975, Río LaNovia,189m,19Feb2002, Ponacillo, 570m,26Jun1984, Huinguillo, 500m,05Mar1962, Pasco: Oxapampa,1,850m,09Dec1982, Mar 1976, Tabasco: Balancan,LaPalma,01Jun1939, Oaxaca: Comaltepec,Tuxtepec, 1,100m,25Jun1966, Soteapan, 29Jun1982, 1004, 1038,1058 (MO*); Catemaco,UniversityofTexas greenhouse, 25May2005, Matuda1404 1974, 12 Aug 17 Mar1983, Feb 1980, 5282 1983, Stevens21532 May 2001, 11651 (MO); Esteli:ReservaNaturalMiraflor, 1,200m,10Jul1999, 1915, Biología LosTuxtlas, 29Mar1979, 3966 (NY); EstaciónBiologíaLosTuxtlas, 28Dec1969, Apr 1969, 1,500 ft,20Jun1978, 3,000 ft,04Jan1968, 1,200–1,500 m,01Jan1961, 16 Jan1979, PERÚ. Amazonas: Bagua, Yamayakat, 320 m,16Jul1994, TRINIDAD &TOBAGO. Andrew Trace: Arima Pass, Universityof VENEZUELA. Anzoátegui: NEofBergantín, 500m,20Feb1945, MEXICO. Chiapas:Berriozábal,900m,21 Apr 1972, SURINAME. Brokopondo: Sectie,19Oct1944, PANAMA. BocasdelToro: WofCampamentoCorrienteGrande,23 NICARAGUA. Boaco:NEdeMombachito,11 May1982, (MO);Zelaya:Siuna,300–400m,15Mar1984, (F*);Hidalgotitlán,140m,07Sep1974, (MO);Jinotega:Wiwilí, ReservaNaturalKilambé,900–1,200m,24 (MO*);Huambisa,180m,23Nov1979, (US); Atures: Territorio Federal Amazonas, 80–100m,04Mar (US);Tambopata Reserve, 250m,09Mar1981, Purpus 7563 Steele etal.1054 Steyermark 89251 Stevens &Moreno 22139 Guanchez 1585 Vásquez &Jaramillo1912 Correa etal.3725 Rueda etal.16360 Lewis etal.5283 Revilla 313 (US*);Tapana Trace: Valencia-Matura Road,30Mar1959, Steele &Meza1052 (MO);Catemaco,0m,13Jul1983, (MO); Lara:Guárico,SanIsidro, Mar1939, Hamilton &Stockwell3454 (MO);NWslopeofCerro ElPicacho,1,420–1,520m,25May Conrad &2870 Dillon &Turner 1416 Stevens 11548 (MO);LosSantos:LomaPrieta,2,400–2,800ft,08Jun1967, (US);Panamá:,Cerro Azul, 24Mar1969, (NY*);GuanapoRoad,29May1975, (MO);Gamboa:CerradoPelado,30–200m,18Mar1983, (TEX*);LasChoapas,160m,06Mar1978, (MO). (F); Aragua: RanchoGrande,02 Apr 1926, (MO*);Trinitaria, 20Sep1984, (MO); Bolívar: NofLasNieves,600–650 m,05Feb (MO*);Maynas,140m,21Nov1991, (TEX*);Madre deDios:Manú, 24Nov1965, (US);Dept.unknown: BalandraBay, 08Mar1920, Duke &Lallathin15007 Calzada 9801 Nee etal.24704 (NY); LaGranSabana,900–1,000m,25 Apr 1988, Hammel 3516 Schunke 1597 Smith 10077 (MO);Darién:SofElReal,50m,31Mar1985, (MO);MacizosdePeñasBlancas,1,200–1,330m, Schunke 9364 (MO);Chiriquí:RíoColorado, 1,200–1,400m, (MO);Matagalpa:SantaMaríadeOstuma, Heller 7 Davis etal.770 (MO*);SanJuan:0m,27Mar1971, Knapp &Mallet6542 (TEX*);RíoMomon,Nanay, 125m,21Jan Steele 1056 (MO);Maynas,Iquitos,RíoMomón,09 Dillon etal.1780 (F);Loreto: Alto Amazonas, 210m,05 Schunke &GrahamS14849 Thieme 186 (F);FuentePura,1,400m,05Jun1982, (MO);Veracruz: Atoyac, 13May1937, Woytkowski 7150 (F);Palenque,170m,11 May1982, (F*);Junin:51kmNEofTarma, 1,550m, (MO*);Colon:SantaRitaRidge,06 Schunke 5804 Martínez 12472 (US*); Arima: 04Mar1921, (NY);Tuxtla, 17Jun1971, Hawkins &Mejía504 (MO);Coclé:Cerro Pilon,ElValle, (MO*);BosqueNacionaldeIparia, Kujikat 219 Croat 19325 Mori etal.4680 Smith 2916 Izaquirre 167 (F*); Carretera Iquitos,133m, Matuda 3313 Dorantes 3510 (TEX*);SantaIsabella trail, (US). (MO);Continental Divide, Gentry &Dwyer3412 Antonio 2493 Torres &Hernández3281 (MO*);LeoncioPrado, Irwin etal.54907 (TEX*);Zacualpan,Dec (MO*);Ucayali:Purús, (MO);Huánco:Colpa, Tunqui 147 Maguire &Stahel25003 (MO*);Ocozocoautla, Ortiz 1889 Gómez-Pompa &Rosas (MO);Río Ampiyacu, (F);SanMartín,Río Woytkowski 7590 Philcox 7822 (MO*);SanMartín: Méndez 7939 Martínez 906 (MO);LaMuralla Lindeman etal.402 Méndez 186 (NY);50–100m, Tamayo 685 Young 42 Porter etal.4088 (MO); Altos de (MO);Estación Breedlove 24811 García 554 Vásquez 17577 (NY). (MO);Barro (MO);Santa Sandino 2823 Vásquez etal. (MO). Croat 12218 (MO);Río Croat 25652 Rueda etal. Ward 7904 Britton & (NY). (NY). (MO); (TEX); (MO); (MO); (MO); Nelson Vargas Pittier (US); Steele (F*); (F);

and theremainder were sterile. 74.4% hadmale flowers,23.3%hadfemale flowers orfruits, peak Januarythrough June.Ofthe43specimensexamined, wide, 1.5–1.7mmthick. thickness unknown.Seeds30–78, 5.0–6.0mmlong,2.8–3.0 lighter green stripes,21–39mmlong,12.5–18.0wide, wall long, stigma3.5–5.0mmlong. Fruits, green, sometimeswith oblong, 10.0–15.0mmlong,2.3–6.5wide,style4.5–5.5 long, 2.4–2.9mmwide,length-to-widthratio2.1;ovaryelliptic- als orange,oblong-elliptic-rhombic, apexacuminate,4.5–6.5mm ear, colorsameasordarkerthancalyx,1.0–2.0 mmlong;pet- long, 1.3–6.5mmwide,adaxialsurfaceglabrous; sepalsthin,lin- salverform, cylindrical,puberulent, solidgreen, 20.0–27.5mm 0.3–0.8 mmlong.Pistillateflowers2–3pernode,axillary;calyx 4.8–11 mmlong,antherappendageround, minutelypapillose, long, 2.5–7.3mmwide,length-to-widthratio2.0;anthersstraight, als orange,ovate-elliptic-rhombic, apexacuminate,5.5–16.0mm sepals thin,linear, colordarkerthancalyx,1.0–3.5 mmlong;pet- 6.5 mmwide,adaxialsurfaceglabrous withtrichomesatthroat; light green withdarkerspeckles, 5.5–12.0(–21.0)mmlong,1.5– than subsequentflowers;calyxsalverformflask,puberulent, ers permm.Staminateflowers,first2–3flowersnotmuchlarger pedicels puberulent, (0.5–)1.0–19mmlong,with0.24–0.7flow- with darkersplotches,0.5–1.6mmindiameter, 70–205mmlong; corymb; pedunclesglabrous orrarely puberulent, lightgreen average 73.8%oftotallength.Inflorescence anindeterminate acuminate, 55–180mmlong,(23–)31–70wide,lobedepth 3– to5–lobed,baseauriculate,(5–)10–33(-40)mmdeep,apex glabrous, 0.7–2.0mmindiameter, 12–75mmlong;simpleleaves extending beyondblade,simple;petiolespuberulent orrarely main veins,flush,margins entire, withirregularly-spaced veins nous, bothsurfacesglabrous withveinsseldompuberulent, 3–5 with darker, speckles,0.2–1.5mmindiameter. Leavesmembra- eter; tendrilsglabrous orpuberulent, solidgreen orlightgreen Anguriawarmingiana Anguriaintegrifolia 1980, Diederichs72 1979, 119531 Tachira: MontañadeGuafitas,250–300m,07Nov1979, Stannard&Liesner339 (MO); RíoNegro: Territorio Federal Amazonas, 1,250m,23Mar1984, 107 Anguriajacquiniana 5. Jan 1922, 29 Aug 1973, Morán, 05Mar1983, Phenology— Stems glabrous orpuberulent, solidgreen, 1.2–3.9 mmindiam- Psiguria umbrosa (TEX);Portuguesa:Sucre, 1,550m,16Mar1985, online photographBR!). 354. 1978.—TYPE:BRAZIL.1860, Psiguriawarmingiana Dec 1816,W XII. 1.9.1825.—TYPE:BRAZIL.S.Pedro d’Alcantara, 242–246 442. 1973.—TYPE:N.J.Jacquin,Sel.Stirp. Amer. Hist. quiniana photograph: F!). Bordones, 2:121. 1817.—TYPE:VENEZUELA.Sucre: Camanacoa, 347. 1978. Steyermark etal.123263 Davidse &Gonzáles16553 (MO);Trujillo: 900m, Pittier 10142 (NY*);Zulia:Miranda,Cerro LosManantiales, 600m,03Jun Tillett &Hönig738–544 (Schltdl.)R. A. Howard, J. Arnold Arbor. 54(4): tabula 156 Flowering throughout theyearwithflowering Bonpland 165 Anguria umbrosa ied s.n Aymard etal.1889 (US);GuatopoNationalPark,20Feb1981, (F);RíoBaria,140m,17Feb1985, NeesetMart.Nov. Acta Acad. nat.cur. Schltdl.,Linnaea24:708.1851. Cogn.Diagn.Cucurb.I:21.1876. (Kunth) C.Jeffrey, KewBulletin33(2): . 1763.(holotype!) . (holotype:onlinephotographBR!). (MO*). (Cogn.)C.Jeffrey, KewBull.33(2): Pittier 13522 (MO); Yaracuy: El Amparo, 08Mar1973, , (holotype:P fide Jeffrey (1978) ; (NY);Miranda:ElPortachuelo,29 (NY);Mérida:Tovar, 600–800m, Kunth,Nov. Gen.Spec. (F*);Tucupita: 200m,04 Apr Peckolt 512 Ortega &Smith2575 Nee 30916 Steyermark etal. Psiguria jac- (holotype: Condon (NY);

Delivered by Publishing Technology to: University of Missouri-Columbia IP: 128.206.9.138 on: Wed, 17 Aug 2011 13:14:52 Copyright (c) American Society for Plant Taxonomists. All rights reserved. lengths ofpedicelsonmaleflowers( cation. Themostconsistentcharactersincludetheaverage but variationinthesecharacteristicscanconfoundidentifi- lobe; speckles, andpetalsare light orange.In 4.3 mm),pedunclesandcalicesare lightgreen with darker but there are simple,trilobedleafmorphsof from thesimple,3–to5–lobedleavesof the trifoliolateleafmorphologyof found underdescriptionsofthosetaxaorthekeys.Typically, distinguish most edgeofthe comes intocontactwith part ofthebasemap image. trilobed leaves( zii and apex( [Volume P. 35 warscewiczii rosa 2010). However, iffoundinthesmallregion ofoverlap, P. warscewiczii supported cladeof low base. are solidgreen, andpetalsare deepred, typicallywithayel- rescences are spicate(pedicelsabsent),pedunclesandcalices cences of flowers are neededtodistinguishthespecies.Maleinflores- 5K–O ) thatcanconfuseidentification.Inthoseindividuals, SYSTEMATIC BOTANY laps with shrubs, orfallentrees. ravines, inshadeoralongforest edges,climbingoverrocks, moist tropical forests), alongroadsides, riverbanks,slopes,or exclusively intropical dryforests (withfewindividuals in ern Brazil( Fig. 13 ). Foundatelevationsfrom 15–850m,almost America andtheLesser Antilles, with 354 Fig. =darkorange,rounded-acute), andtheoutermargins of The molecularphylogenyof Discussion— Habitat— and Distribution canbedifficult todistinguishfrom thetrilobedmorphof P. warscewiczii 13 . . P. umbrosa P. racemosa, P. ternata Geographic distributionof P. umbrosa P. umbrosa . Differences mayexistsuchasmalepetalcolor with moderatebootstrapsupport(Steeleetal. The distributionof P. umbrosa P. warscewiczii =larger indentationandlaterallobe), are racemose(pedicelaveragelength= =lightorange,acuminate; P. umbrosa rm from P. warscewiczii =smallindentationandlateral P. racemosa Predominantly innorthernSouth , and range.Characteristicsusedto Psiguria samplesthatwassisterto Psiguria umbrosa P. triphylla P. triphylla a fewindividuals Psiguria umbrosa P. umbrosa resulted inastrongly P. triphylla P. umbrosa n and in thesoutheastern- P. ternata ifrnits it differentiates P. triphylla . Opencircles are ad possibly and , P. warscewic- i. T U), 5T, (Fig. ml inflo- male , =4.3mm; a be can in east- P. umb- over- (Fig. thin andlinear; 1937, Irwinetal.18941 10 Apr 1966, 8512 (NY*); Pará:Concei Crato, 700m,19Jan1983, asterisk were measured forspeciesdescriptions) P. warscewiczii Anguriaovata 6. 62797 Fernández4014 Hierro, 21 Aug 1961, Monagas: Caripe,850m,17 Apr 1945, 3789 1903, 01 Nov1962, Mamo ElectricPlant,23Jun1923, Broadway 226 10 Mar1911, Hacienda deCura,480–1,200m,05Jul1918, Pittier6048 1982, Maracas Falls,650ft,30May1975, 2207 Coker&Rowland658 Juan, 13Jun1903, Old Province Road,20Jun1981, Acevedo 3302 ulent, veinsflush orseldomabaxialandadaxial mainveins both surfacesglabrous withveinsandmargins seldompuber- diameter. Leaves immature membranous, mature coriaceous, puberulent, lightgreen withdarker, speckles,0.4–1.8 mmin ear splotches,1.5–5.0mmin diameter;tendrilsglabrous or Anguriaaurantiaca Anguriatonduzii Anguriamagdalenae Angurialimonensis Stems glabrous orpuberulent, lightgreen withdarker, lin- Psiguria warscewiczii TRINIDAD &TOBAGO. Tobago: Great DogRiverValley, 12Oct FRENCH GUIANA. Mt.Rorota, ancientvolcano,24 Apr 1992, BRAZIL. BarradoCorda: 08Mar1983, Examined— Specimens Representative ST. VINCENT. 500–1,000ft,15Oct1890, ST. LUCIA.Barre del’Isletrail,24Jan1985, MARTINIQUE. 1880, VENEZUELA. Aragua: LaTrinidad deMaracay, 440m,Feb1913, (F*,MO*);Paraíba:24Jun1959, (MO*);Cúpira,0–150m,16May1981, (NY); Arima Valley, 1325ft,12Jun1973, 38(3): 219.1978. (holotype: onlinephotographK!). 88 t.5304.1862.—TYPE:PANAMA. May1861, Paraíso, 12May1919, HONDURAS. Copán:trailfrom HaciendaElLimónto El C. Nelson,Fern.&Casas,Fontqueria52:4.1998.—TYPE: 24(1): 26–27.1922. (1978) ; onlinephotograph!;isotype:US!). Dulce, Apr 1896, 1916.—TYPE: COSTA RICA.SantoDomingodeGolfo foothills, 4Jul1906, Río FríobetweentheCiénegadeSantaMartaand Fig. 24.1910.—TYPE:COLOMBIA.Magdalena:around Moin, Sep1899, 25. 1910.—TYPE:COSTA RICA.betweenPortLimonand 13006 COSTA RICA.LasVueltas: Tucurrique, Feb1899, (Donn. Sm.)C.Jeffrey, KewBull. 33(2):353.1978.—TYPE: (F);Ríoelmedio,200m,15Jul1972, Harriman 17631 Johnston 287 Sandwith 1748 (US);Villa deCura,03Dec1962, (holotype:US!). (NY). Irwin etal.14593 (NY*); Aricagua, Feb1923, Steyermark 90976 Bond etal.243 (NY);RíoGuagua,230–300m,18May1945, Donn.Sm.Bot.Gaz.31:112. 1901. =1.0mm)andsepalcharacters( (F*);Maranhão:Fortuna,21Feb1983, (GH);Miranda:Brión,0–30m,27May1981, P. warscewiczii ç Johnston 14 (NY);Plymouth,16Jan1953, Cogn.Pflanzenreich 66(IV. 275.I):191. ão do Araguaia, 350–620m,08Feb1980, (US*);Matchepoorie,11 Mar1921, (NY); Aristequieta 4748 Pittier, Contrib.U.S.Nat.Herb.13:119 Fig. S.F. Blake,Contrib.U.S.Natl.Herb. Pittier, Contrib.U.S.Nat.Herb.13:118 Plowman &Ca Pittier s.n Anguria warscewiczii Duss 750–117 Tonduz 9999 Broadway 2009 (US);CristóbalColón:nearriver, Jan1923, (NY);Corumbá deGoiás,700m,21Jan1968, Pittier 1630 (US*);IslandofMargarita: 450m,12 Aug (GH);HeightsofSt. Ann, 17Mar1920, Psiguria aurantiaca Blake 7355 (Hook. f.)Wunderlin, Phytologia Baksh &Adams481 Pittier 11076 Philcox 7825 =thickandtriangular). . (holotype:US!;isotype:NY!). 2201 (NY);Valdez, Güiria,27May1983, (NY*). Steyermark 62159 ç ula 12742 (US). (F*). (holotype:Kfide Jeffrey (holotype:US!). Dumont etal.VE-7685 Liesner &González11901 (collections markedwithan Broadway 493 (holotype:US!). Schatz etal.890 Smith &Britton234 Trujillo 5493 Pittier 7931 Philcox &Wood 7079 Howard etal.19874 (US*);LaSabana,100m, (NY*);RuizTrace, 21Jan (F*);Goiás:Posse,800m, Hook.f.,Bot.Mag. F. W. H.19999 (NY*);Trinidad: San (S.F. Blake) (F);Sucre: Puerto (GH);Caraquita: (US);Carabobo: Britton & (GH);Federal: Psiguria ovata Santos etal.646 P. umbrosa (S) Ceará: (US*); Plowman etal. Hayes s.n Steyermark (NY). Berry etal. (NY*). Grimes & Tonduz (A*). (GH*); (MO); (NY); = .

Delivered by Publishing Technology to: University of Missouri-Columbia IP: 128.206.9.138 on: Wed, 17 Aug 2011 13:14:52 Copyright (c) American Society for Plant Taxonomists. All rights reserved. many formsas that in Leaf morphology in sion undereachoftheirtaxonomic descriptionsorthekeys. P. warscewiczii P. warscewiczii contact with lection of overlaps withthoseof sandy, orclaysoils. or slopes,climbingoverothervegetation,andonlimestone, found indenseshadeorlightgaps,onroadsides, riverbanks, ary forests, cloudforests, andfloodedgrasslands.Maybe extend intodryareas alongriverbanks),primaryorsecond- and subtropical evergreen orsemideciduous,moist(butcan Found atelevationsfrom sealevelto1,600mintropical Mexico tonorthwesternColombiaandVenezuela ( Fig. 14 ). female flowersorfruits, andtheremainder were sterile. specimens examined,76.8%hadmaleflowers,9.6% 56–88, 6.4–8.7mmlong,3.8–4.8wide,1.9–2.7thick. 34–80 mmlong,14–22wide,wall0.7–2.2thick.Seeds 7.2 mmlong.Fruits, green withlighter green orwhitestripes, mm long,1.9–3.5wide,style2.3–4.5stigma5.5– wide, length-to-widthratio1.3;ovaryelliptic-oblong,10.0–15.0 elliptic-orbicular, apexcuspidate,4.5–9.2mmlong,3.3–6.9 than calyx,0.5–1.6mmlong;petalsbrightorangeororange-red, trichomes atthroat; sepals thick,triangular-linear, colordarker long, 1.0–3.6mmwide,adaxialsurfacepuberulent withsparse drical, glabrous orpuberulent, striped green, (4.9–)18.0–25.0mm ends ofbrancheswithreduced leaves;calyxsalverform,cylin- mmlong.Pistillateflowers2pernode,axillary,(–0.7) butatthe or rarely triangular, glabrous orrarely papillose,0.1–0.5 straight, (1.0–)6.0–10.9mmlong,antherappendageround long, 3.0–7.0mmwide,length-to-widthratio1.6;anthers ovate-elliptic-orbicular, apexrounded-acute, 4.0–10.0mm calyx, 0.5–2.1mmlong;petalsintenseorangeororange-red, trichomes atthroat; sepals thick,triangular, colordarkerthan mm wide,adaxialsurfaceglabrous orpuberulent withsparse light green withdarkerspeckles, 7.0–17.0mmlong,1.0–4.6 355 calyx salverform,cylindrical,rarely flask-shpaed,glabrous, first 2–3flowersnotmuchlarger thansubsequentflowers; long, with0.25–0.75(–1.2)flowerspermm.Staminateflowers, (36–)130–285 mmlong;pedicelspuberulent, (0.1–)0.5–1.9mm light green withdarkersplotches,1.0–1.9mmindiameter, long. Inflorescence acompactraceme;pedunclesglabrous, attenuate andabaxialbaseacute-truncate, 40–115(–145) mm leaflets onpedateleavesbaseoblique,adaxialacute- acute-truncate, 25–169mmlong,14–69wide;outer base oblique,adaxialacute-attenuateandabaxial ate, 52–206mmlong,17–75(–150)wide;sideleaflets large, apicesacuminate;centerleafletbaseacute-attenu- STEELE:TAXONOMIC REVISIONOFPSIGURIA and sideleafletsovate-ellipticwithlaterallobe,sometimes plex leaveswithcenterleafletelliptic-rhombic-oblanceolate ulent, 0.6–1.5(–2.1)mmindiameter, 0–20mmlong;com- average 73.5%oftotallength;petioluleswingedandpuber- nate, 86–183mmlong,30–90(–122)wide,lobedepth lobed, baseauriculate,10–25mmdeep,apexacumi- 0.8–2.7 mmindiameter, 12–96mmlong;simpleleavestri- or seldompedate;petiolespuberulent orrarely glabrous, spaced veinsextendingbeyondblade,simple,trifoliolate, prominent, margins entire-irregular-lobed, withirregularly- 2010] Discussion— Habitat— and Distribution Phenology— P. pedata P. umbrosa rm from range. Forcharacteristicsusedtodistinguish Flowering throughout theyear. Ofthe250 The geographicrangeof (inCostaRica),andpossiblycomesinto P. racemosa P. pedata P. warscewiczii inthesoutheastern-most edgeofthe P. racemosa,P. triphylla ; therefore, thebestwaytodistin- n and otnna fo southern from Continental ( Fig. 5V–BB ) cantakeas P. umbrosa Psiguria warscewiczii , seethe discus- , andonecol-

support asplitofthesecladesintotwospeciesinthefuture. tional molecularand/ormorphologicalcharactersmay graphic lines(anortherncladeandasouthernclade),addi- clades, sincetheinnercladessplitpredominantly alonggeo- et al.2010). Although thetrifoliolatemorphis foundinboth this cladewassplitintotwoweaklysupportedclades(Steele ported, monophyleticcladeof 5V–BB ). Themolecular phylogenyrevealed aweaklysup- gins andlaterallobesizes(foundthroughout therange)( Fig. end oftherange),andtrifoliolateleaveswithvariablemar- ern endoftherange),pedateleaves(foundatnorthern simple, trilobedleavesofvaryingdepths(foundinthesouth- flowers permm). ( and calyxcolorshape( age pedicellength( ( guish thetwoiswithmaleflowers.Theydiffer inanthershape Marta: 300ft,30sep 1898, 2170 1943, Jul 1975, Palmar, 450m,30 Aug 1976, (NY); Riosucio,16Jun1957, 2139 Smith 14260 Mar 1949, 1979, Salgar, 1,410m,11 Nov1989, 4954 Jan 1994, Machaca, 50ft,07 Aug 1933, ( solid green) andthedensity offlowersalongthepeduncle zii green, oval).Thebestcharacters todistinguish darker speckles,cylinderorelongatedflask; are partofthebasemapimage. asterisk were measured forspeciesdescriptions) P. warscewiczii P. warscewiczii P. warscewiczii Morphological variationin rm from Fig. COLOMBIA. Antioquia: Arboletes, 30m,23Mar1987, BELIZE. StannCreek: Middlesex,24Jul 1939, Examined— Specimens Representative (MO);RíoSanJuan: 5m,31May1946, (US);Chocó:BahíaSolano,0–75m,22Feb1939, (MO); Sabanalarga, 1,100m,08 Apr 1986, Renteria 1731 Haught 3851 14 Gentry &Enrique15301 . . Lowman &Foster74 von Sneiderns.n P. triphylla (NY*);Caldas:La Dorada, 200–400m,30Dec1936, Geographic distributionof =lightgreen withdarkerspeckles; =straight; = (NY*);Mutatá:150–200 m,05May1987, (MO);Bolívar:Quimarí, Cordillera Occidental, 500m, < P. warscewiczii 0.75flowerspermm; includestem,peduncle,andcalyxcolor . (NY*);Torrecilla, 150–300m,07Nov1926, Smith 1601 Forero etal.2364 Girón &Ortíz263 Schipp S-298 (F). Romero-Castañeda 6263 P. pedata (MO); Magdalena:Codazzi,100m,17Nov P. warscewiczii (F, NY);Santander: Cimitarra,200m, Psiguria warscewiczii P. warscewiczii =1.0mm; Psiguria warscewiczii =foldedbackwards), aver- (GH);BlueCreek, 100–200 m,07 (collections markedwithan (MO);Unguía,100–300 m,19 (NY*); Segovia,750m,21Jul Cuatrecasas 21527 Callejas etal.2271 Gentle 2937 P. triphylla =lightgreen with P. pedata (GH);SanJosédel Killip &Garcia 33467 P. pedata ape, and samples, P. warscewic- P. triphylla Fonnegra etal. Zarucchi etal. . Opencircles =5.2mm), (F);Toledo: includes = (F);Santa solid = Killip & > (NY*); Haught 0.75 =

Delivered by Publishing Technology to: University of Missouri-Columbia IP: 128.206.9.138 on: Wed, 17 Aug 2011 13:14:52 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 1969, Luis, 09Oct1966, Oct 1911, Prieta, 800–900m, 08Jun1967, Santa Fé,30–50m, 26 Apr 1982, Punta GuayaboGrande,0–200m,25 Jan1982, 1979, Río Pirre, 14Jul1971, Contreras 35–150 m,07 Apr 1940, Puerto Viejo, 45m,12 Aug 1972, (F); CantóndeLiberia,750m,13 Aug 1996, 1964, 1905, Escoba, 0m,03May1939, 23488 Steele&Fernández1006 12 Jun1996, Corcovado NationalPark,0–5m,06Jul1977, (MO); Puntarenas: OsaPeninsula,100ft,12 Aug 1967, (MO); Cordillera deTalamanca, 400m,03Sep1988, Croat 522 300–500 m,12Mar1924, Research Station,100m,02 Apr 1980, Real, 06Jul1962, Jun 1959, 300–400 m,12Dec1981, Gra 2237 Zone, 08Oct1939, Llano Grande,450m,07May1981, Alston8788 5 SSEAI OAY [Volume 35 Quirigúa, 72–150m,26 Apr 1939, 1996, SYSTEMATIC BOTANY 11 Sep1995, de Golfito,50m,14Sep1992, Quebrada Bonita,35–40m,11 Jun1986, 700 m,27Jan1926, Turrialba, 500–600m,21Jul1949, (MO); RanchoGrande,500m,14Jul1991, H1691 4,800 ft,24Jul1962, 27 Jul1975, 356 Bárbara: Nueva Arcadia, 13 Aug 1970, (MO*); Lempira:Cuábanos,1,600m,25Sep1963, Progreso, 12 Aug 1929, Tela River, 03 Apr 1903, unknown: LasVegas, 3,500ft,21Jun1970, Nelsonetal.2841 Montañuelas, 1,400m,18Jul1962, 100–150 m,08Jun1985, 3433 1982, 1980, (MO*). Nov 1988, Mombacho, 600m,02Jul1983, Boaco: ElPortón,370m,16Jul1978, Ana, 210m,06Dec1950, San JuanLindo,200m,01Dec1950, Molina26214 27 Jan1976, Sytsma&Andersson4566 10 Aug 1974, Burica Peninsula,40m,06Mar1973, 446 28122 10434 1985, 02 Aug 1980, 21 Aug 1984, Cedro, 700m,19Jun1980, (F);ChiriquiLagoon,09May1941, GUATEMALA. Alta Verapaz: Panzós,280m,04Sep1988, HONDURAS. Atlántida: Tela, 25ft,21Mar1926, COSTA RICA. Alajuela: MuelleSanCarlos,975m,01Mar1939, NICARAGUA. Atlántico Norte:Cerro Livico, 400–600m,12Dec MEXICO. Chiapas:Ocosingo,570m,04Oct1976, PANAMA. BocasdelToro: Changuinola Valley, 22Feb1924, (MO);NuevaGuinea,300m,13 Aug 1982, Contreras 8809 (TEX);Tucuru, 280m,04Sep1988, (MO). (MO*);Rivas:Volcán Maderas,600–800m,15Sep1983, Moreno &Sandino14957 Stevens 18708 Hammel etal.20374 Deam 5 Moreno 25513 Williams etal.26451 Hammel 7313 (F);SanRamón,1,100m,25 Apr 1983, (US);Cerro Bruja, 20May1978, 11117 (MO);Cerro Coronel, 20–170m,15Sep1986, Stern etal.134A Pittier 4541 MacDougal etal.3293 (US);Penonomé,1,200 ft,24May1967, Gentry &Forero 15477A Sanders etal.17856 Molina etal.31471 Croat 26708 (F);Copán,16Jun1977, (GH);Quirigúa,75–225m,May1922, Cascante 1018 Moreno 1662 Moreno 24467 (LL);Peten:Dolores, 21Jun1961, (MO);ElCacao,600m,25Dec1982, Duke 5146 Contreras 6339 (MO); Atlántico Sur:CañoMontecristo,100m,06Feb (MO);Manene,22Dec 1980, (MO);Castillo,100–200m,17Mar1999, (F, LL*). Standley &Valerio 46213 Robinson 97 Allen 2011 (US);MatíasHernández, 25 Aug 1914, Croat &Porter15539 (TEX*). (MO);Coclé:ElCopé,1,200–1,300m,13May1931, Bangham 365 (MO*);SanJosé:CantóndeMora,900–1,000m, (MO);Paya,12Jun1959, (MO);ElValle de Anton, 1,000m,05Jun1939, Wilson 658 Standley 37300 Knapp &Sytsma2404 Moreno 872 (F);Tilarán, 03Jan1964, Steyermark 39024 (MO);RíoSanJuan:La Palma,50m,21Mar (F);CiudadColón,900–1,000m,20Jul2005, Molina 3611 (MO); NuevaSegovia:ElJícaro, 550–600m, (MO);Pucro, 22Jun 1967, Téllez 8760 Aquilar 1305 (MO);Zelaya,70–80m,31Jul1982, Standley 72877 (TEX);CantóndeGarabito,20–30m,17 Aug (F);Copán:SantaRita,700m,21 Aug 1971, Duke 11851 (MO);Portobelo,30Jan1973, (MO). (US). (LL*);Dept.unknown:LaCumbre, 31Jul Grijalva &Ayesta 2722 Opler 1608 Huft etal.1969 Holm &Iltis438 Standley 72209 (MO);Valle delCauca:NWofDarien, (NY); Yoro: SanJosé,200–600m,06 Sytsma etal.4426 Molina 10864 Croat 22583 Hammel 8408 Stevens 9265 (MO);Matagalpa:Ranchería,280m, (A);Jardín BotánicodeLancetilla, Molina 3479 Harmon &Dwyer3809 Poole &Watson 1167 (F);PuertoCortés,10 Aug 1975, (US); Airport Limón,26Jul1965, (TEX);Comayagua:Quebrada Grayum etal.7601 (MO); Panamá:Chepo,60–80m, (MO);Garachiné, 500ft,09May (MO);NicoyaPeninsula,200m, von Wedel 2429 Hammel 3218 (MO);LaSelvaOTSBiological (F);ElEstor, 03Mar1972, Martínez 23490 Barkley &Smith40860 (US);LaCruz, 200m,01Jan (F);PuertoBarrios,25Feb Morales 5639 Rivera 1449 Knapp &Mallet3153 Liesner &Judziewicz14894 (MO);Darién: Yaviza, 06 (MO);SanBartolo,125m, (MO);LosSantos:Loma Liesner 2893,3001 (A);Guanacaste:Serena, Contreras 2497 Molina 12904 (F);MontañadelMico, (MO);Granada:Volcán (MO);Limón:Guápiles, (NY);SanJosé,300m, Araquistain 3095 Stern etal.414 Hartman 12122 (GH);MontanaSanta Lewis etal.1521 Standley 24208 Williams etal.26563 (MO);Colón:Canal Albertina etal.32001 Raven 21722a (MO);Jinotega:El Grayum etal.8730 (MO);Salamanca, Calzada etal.2675 Duke 13072 Mitchell 37 (GH);Chiriqui: (MO);Cartago: (NY);Heredia: (MO);Cantón (MO);Izabal: (LL*);Cortés: Stevens 24587 (MO);Dept. Nee &Téllez (US);Santa Pittier 6765 Rueda etal. Kennedy & (LL*);San (MO);El Martínez (MO*); Sandino (GH*); Dunlap (MO); (MO); (MO); (MO); (MO); (GH); (US); (NY); Smith

A.triphylla Anguriarosea Anguriaplurilobata Anguriaparviflora & Miller1001 Pedro Miguel,17Jun1938, Island, eastoflaboratory, 23Dec1931, (US); Barro ColoradoIsland,12Jul1927, 08 Aug 1982, Gigante Peninsula,27Jul1982, 1977, Cno, . . 94. 1984. A. M. Condon, Melothrieae. & Cucurbitaceae-Fevilleae 1916. A. Cogniaux, Anguriaaffinis Alcalde Diaz,190m,23Nov1973, Croat 8499 1967, 2,000 ft,26Nov1966, (MO); FortClayton,25May1966, Oct 1961, Bartlett&Lasser16336 Perija, 300–700m,25 Aug 1967, 2340 08 Sep1982, (MO); Santiago,150m,17Jul1976, 7699 Cgiu, . 81. Ccriaés. p 6 7 n in 663–679 Pp. Cucurbitacées. 1881. A. Cogniaux, exploi- pollen of Patterns 1981. Gilbert. E. L. Cucurbitaceae. and Smiley , T. the J. L., C. On Boggs, 1841. W. A. G. Arnott, New York Botanical Garden forsupplyingtheNeotropical base map. the work,andWayt Thomas,Ph.D.,InstituteofSystematic Botany, The ing facilitiesandstaff assistance thatmaintainedlivingcollectionsusedin collecting permits,UT Austin’s BrackenridgeFieldLaboratoryforprovid- M. Timaná (PontificiaUniversidadCatólica delPerú) forhelpwithplant- Bolivia), andK.Meza(Perú) forfieldassistanceandcollectingplants, Botánico NacionalDominicanRepublic),O.Plata(Herbario de Rica), F. Axelrod (UniversityofPuertoRico-RíoPiedras),T. Clase(Jardín and TEXherbariaforloanedspecimens,R.FernándezP. Protti (Costa and theBotanicalSocietyof America. IalsothankMO,NY, F, GH,US, The UniversityofTexas at Austin, the American SocietyofPlantTaxonomy, Dissertation Improvement GrantDEB0808294,thePlantBiologyProgram at The authoracknowledgesthefollowingresearch grants:NSFDoctoral Simpson, L.E.Gilbert,C.R.Linder, andM.Mehdyonearlierversions. ymous reviewers forvaluablecommentsonthemanuscript,andB. Cgiu, . 86. Genre 1876. A. Cogniaux, Acknowledgments. VENEZUELA. Zulia:Perijá,17Oct1966, (US). (MO);Veraguas: SantaFé,500–1,000m,12Dec1971, of Neotropical vines 178–230. 275.I): (IV., 66 Phaenerogamarum tation by Botanique Tome XXVII. bitacées nouvellesetobservationssur lesespècescritiques 1848. nonMiq.Type unknown. 1916.—TYPE: CostaRica. type: B,destroyed). 1916.—TYPE: ECUADOR.Jan1897, destroyed). BRAZIL. Type unknown. destroyed). var. 10. 1876.—TYPE: ANTILLES. Burch 3389 Folsom 3517 (MO*);Carti-Tupile, 200m,18Oct1972, Duke 4485 affinis Hamilton etal.1238 Klotzsch,Schomb.ReisenBrit.GuianaIIIp.88. (MO);SanBlas:Cangandi,10m,05 Apr 1986, : 271–280. 3: Heliconius Hamilton 519 Kunth,Nov. Gen.Spec.2:122.1817.—TYPE: (MO);Barro ColoradoIsland,Lathrop Trail, 09Mar1969, Schltdl.,Linnaea24:760.1851. Buels: F Hayez. F. Bruxelles : (MO);Cerro Cabra,30May1978, (Schltdl.)Cogn.,Diagn.Cucurb.I:22,27: (GH);Tocumen airport,08Sep1963, Cogn.,Pflanzenreich 66(IV, 275,I):190. Reproductive biology, demography, andnatural history . o. 3. Prs: G Masson. G. Paris : 3. Vol. . Cogn.,Pflanzenreich 66.(IV. 275.I):191, Dwyer 7112 (LL);RíoChagres, 1miNofMaddenLake,07 Btefis. Butterflies. Gurania The authorthanksR.K.Jansenandtwoanon- Anguria xldd Taxa Excluded ieaue Cited Literature (MO);Cerro Campana,03 Aug 1983, White 133 (MO);IslaCoiba,03Sep1995, Steyermark &Fernández99719 and Schmalzel 726 (MO);Cerro Brewster, 1,000ft,14Dec Blum &Tyson 2328 ut. p 5 i in 9–15 Pp. Auct. Nee 8304 Sullivan 421 Oecologia Psiguria Wetmore &Abbe32 (MO);Lefevre Park,28Jun1940, Tonduz 8175 Kenoyer 567 Kunth s.n (US);BayanoGuipo,07Jun de Bruijn1226 8 284–289. 48: (Guraniinae):astudy ofthe (MO);Piriatí,200–400m, (MO);SantaFé,1,300m, Eggers 15496 Hooker’s LeJournalde (US*);Barro Colorado Candolle Monographiae Kennedy 1789B Hammel 3265 (MO);GoofyLake, Diagnoses decucur- (holotype:B, . (holotype:B, Anguria pedata (F*);CanalZone, (US);Sierrade Dwyer 4369A (NY). Pflanzenreich Gentry 3010 . Aranda etal. Nevers etal. Fascicle1 , (holo- , (MO); (MO); Miller .

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