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LECTOTYPIFICATION OF PASSIFLORA AFFINIS (PASSIFLORACEAE) AND DISCUSSION OF ITS GEOGRAPHIC RANGE WITHIN THE UNITED STATES PDF

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LECTOTYPIFICATIONOFPASSIFWRAAFFINIS (PASSIFLORACEAE)ANDDISCUSSIONOFITSGEOGRAPHIC RANGEWITHINTHEUNITEDSTATES DouglasH.Goldman HarvardUniversityt-ierbaria 22DivinityAvenue Cambridge,Massacliusetts02138,USA. d(][email protected] ABSTRACT GeorgeFngelinann.inhisdescriptionofPassijloraajjinis.neverdesignatedalioloi)peforthespe- cies,norhasalectotypebeendesignatedinsubsequenthteratureaddressingHngehnann'sworkor NorthAmericanPassijlora.Alectoypeisdesignatedhereandadiscussionandclarificationolthe geographicrangeofthisspeciesintheUnitedStatesisprovided. KeyWords:Passifioraaffmis.lectotypification,geographicdistribution.FerdinandLindheimer, GeorgeEngehnann,CharlesWright,CharlesParry,BennoMatthes RESUMF.N GeorgeEngelmann,ensudescripciondePassijloraajjinis.nuncadesignounholotipoparalaespecie, nitampocosehadesignadounlectotipoenlaliteraturasubsiguientec|uetrataeltrabajodeEngel- mannolasPcis.si/l()?'(:jnorteamericanas.Enestetrabajosedesignaunlectotipo,yademassehaceuna discusionyciarificacidndeladistribuciongeograficadelaespecieenlosFstadosLJnidos. INTRODUCTION PassijloraaffinisEngelm.isawarm-temperateandsubtropicalspeciesoloak- junipersavannasandmesicwoodlandsfromcentralTexasandnortheastern Mexico(Schulz1922;Killip1938;Correll&Johnston1970;Diggsetal.1999). ThetypecollectionwasmadebyFerdinandLindheimerinAugusttoSeptem- ber1849atComancheSpring,BexarCounty,Texas.Thetypeserieswasdistrib- utedin1907bytheMissouriBotanicalGardenasapartofalong-forgottenpor- tionoftheFloraTexanaExsiccata(Blankinship1907),althoughitwasoriginally describedseveraldecadesearlier(Engelmann1850a). Thelabelsdistributedwiththetypeseriesin1907(Fig.1)arepotentially misleading,presentingthetypelocalityas"ComancheSpring;NewBrauntels, etc.,"ComancheSpringandNewBraunfelsoccurringperhaps30milesapart. However,Engelmann(1850b),Blankinship(1907),Killip(1938)andLindheimer correspondencewithGeorgeEngelmann,foundinthearchivesoftheMissouri BotanicalGarden,makeclearthatitwascollectedatComancheSpringalone. Lindheimerspentthemajorityofthegrowingseasonof1849atthatsite,re- turningtoNewBraunfelsonlyattheendofthatyear(Engelmann1850b; SIDA21(1):275-285.2004 276 BRIT.ORG/SIDA21(1) LINDHEIMER, FLORA TEXANA. (supplementaryto "flora texana exsiccata.") Distributed by the Missouri BotanicalGarden. PASSIFLORA AFFINIS, Engelm, TypeCollection. CollectedbyLiii.lheimerfrom1849to1851. Aug.-Sept.,1849. C'omancheSpring;NewBr»«nfels,otc. No. 817. Fig.1.ThecollectionlabelfoundonspecimensofthetypeseriesofPajj/floroflffin/sdistributedin1907(seeBlankinship 1907),thislabelfromaduplicateatGH. Blankinship1907).ComancheSpringwason thepropertyofafriendof Lindheimers,BaronOttfnedHansvon("JohnO.")Meusebach,anearlysettler andfounderof severalcommunitiesincentralTexas(King1967).Thespring wasprobablyveryclosetoMeusebachshouseneartheheadwatersofSalado Creek,inwhatisnowtheCampBulhsMilitaryReservation(Ivey1979). AlthoughEnglemann(1850a)didclearlystatethetypelocalityin the protologue,hedidnotdesignateaholotype,andneitherBlankinship(1907)nor Killip(1938)laterdesignatedalectotypefromtheP.a(finistypeseries. PassifloraaffinisEngelm.,Bost.J.Nat.Hist.6:233.1850.(Fig. 2).Type:TEXAS: [B[-..\ARCo.:]ComancheSpring,Aug-Sep1849,F.Lindheimerl74(lectotyph,here selected:GH!:isoi.ectotypes:ARIZLBM!,BR!,BR1T-SMU(Q,CAM,G!(2),GH(Ki, MFXU,MOlNY(P(PH(TEX!,UQ.US!,W!). Liana,oltensuckeringfromroots,stemsterete,glabrous,minutelypuberulent toscabrouswhenyoung;tendrilsglabrous;stipulesHnear-setaceous,erect,gla- brous,0.1-0.2cm;petiolesglabrous,glandless,0.4-4.5cm;leavesglabrous,en- tire,shallowlytodeeply3(-5)-lobed,broadlyovateingeneraloutline,basally cordate,truncate,tocuneate,l-8(-10)cmlongby1.5-10(-14)ciriwide,lobes ovatetoobovate,roundedtotruncate,entiretoapiculate,oftenbasallynar- rowed,laminarnectariesround,oftenintwoobscurelines,eachrunningfrom baseofleafnearpetioleapextowardthelargersinuses;inflorescenceoftwoor rslairgehltyloyneelo1nlgoawteirngpemrnforduiet,;fflloorwaelrsbrfarcatgsragnlta,berroeucst,,3s,tiopceca+sipoendailcellyd1.e0c-i3d.u5ocums,, greentopurple,Imear-subulatetorarelyovate,0.1-0.3cm;flowers3-5cmin diameter;sepalsmoderatelyreflexed,lanceolatetolinear-lanceolate,rounded GOLDMAN,LECTOTYPIFICATIONOFPASSIFLORAAFFINIS 277 FloraT(^xaiiacxsicata. Bwi-W. Ulils..r.hi,'lh'!m,r,J?/J^\Sm 'liUliil' Fig.2.ThelectotypeoiPassifloraaffinisatGH. 278 BRIT.ORG/SIDA21(1) toacute,palegreentowhite,1.0-1.6cmlong,0.2-0.4cmwide;petalsmoder- atelyre(lexed,linear-lanceolate,subulate,acute,palegreentowhite,0.6-1.3cm long,0.1-0.2cmwide;coronaof2seriesoffilaments,outerserieslinear-fili- form,sinuous,weaklyretlexedornot,taperingdistallyandterminatingina clavateapex,basallypurple,apicallygreen,whiteinbetween,0.9-1.8cmlong, innerseriesfiliform,erect,clavate,palepurplewithgreenbasally,0,15-0.3cm long;floraltubeshallowlyconvextonearlyflat,operculumerect,incurved, plicate,palepurpletowhite,0.1-0.2cmlong,nectarringabsent,limenerect, incurved,white,0.1cm wide;androgynophore0.7-1.0cm long,terete,pale purple;stamenliiamentsgreenwithpurple,0.4-0.6cmlong;anthersgreenwith purplemargins,0.25-0.4cmlong;ovaryglobosetosubovoid,glabrous,styles purple,filiform,glabrous,0.5-0.7cmlong,stigmasbroadlyexpanded,round, 0.1-0.2cmwide;berry,purple-black,ovoid,globose,todorsiventrallycom- pressed,1.0-1.5cmlong,1.0cmwide;seedsdarkbrowntoblack,flattened,obo- void,acute,transverselysulcate,0.25-0.35cmlong,0.2-0.3cmwide,withwhite arillateswellingtooneside;germinationepigeal. ItwouldbeanticipatedthataspecimenchosenasthelectotypeofanEn- gelmannnameattherankofspeciesorbelowwouldbeattheMissouriBotani- calGardenwheremanyofhiscollectionsreside,St.Louis,Missouri,being Engelmann'shomefromtheearly1830suntilhisdeathin1884(White1896; Yatskievych1999).However,theprotologuedescribesbothfloweringandfruit- ingmaterialandnoneoftheexaminedduplicatesofthiscollectionhaveflow- ersandfruitexceptforthespecimenchosenasthelectotypeatGH(Fig.2). Mostduplicatesseenaresterileorhavefewfruit,theonlyotherfloweringspeci- menbeingatBRIT-SMU.Alloftheduplicatesdistributedin1907havemass- producedlabelsnumbered817,whereasoneofthetwoGHspecimens,theone with1lowers,hasamucholder,mostlyhand-written label,numbered 17'^. Lindheimerhadhiscwnifieldnumbers,whereasEnglemannusedseparate numbersbasedonthedistributionorderoftheEloraTexanaExsiccatafollow- ingtheBenthamandHookersequence(Blankinship1907).Blankinshipdoes note(p.170)that174istheLmdheimernumber,whichcorrespondstotheEn- gelmannnumber817,thereforethesenumbersrefertothesamecollection.The handwritingonthelabelofthenow-lectotypeisalmostentirelyEngelmanns, identifyingtheplantasPa'^sifloraIriloba.Elsewhereonthelabelandinthe lowerright-handcornerofthespecimeniswritten"affinis"inwhatisprob- ablyAsaGrayshandwriting.Inaddition,thisistheonlyspecimenofthetype seriesthatindicateshabitatinformationasitiscitedintheprotologue,stating thattheplantwasgrowingin"shadyplaces"and"climbinghighovertrees." Noneoftheseriesdistributedm1907giveshabitatdetails.Theonlydetailin thelabelofthelectotypethatdiffersfromtheprotologueandthe1907labelsis thedate,theolderlabelstating"Sept1849"whereastheprotologueandnewer labelsstate"Aug.-Sept.1849"(Fig.1).However,thisdoesnotmaketheolderla- GOLDMAN,LECTOTYPIfICATIONOFPASSIFLORAAFFINIS 279 belincongruentwiththeprotologue,andbecauseoftheotherinformationon itandthepresenceofflowersonthisspecimenaloneitisthebestchoicelorthe lectotype. GF.OGRAPHICDISTRIBUTION InTexasP.affinisistypicallyioundgrowmgoverCretaceouslimestoneormuch lesscommonlyoverPrecambrianigneousrockorlateTertiarysandstoneand clay(Sellardsetal.1932;Spearmg1991J.InnortheasternMexico(seespecimens examined)itgrowsoverCretaceouslimestone,shaleandsandstone(Direccion GeneraldeGeografiadelTcrritorioNacional1981a,b;PadillaySanchez& Aceves-Quesada1992).ThecurrentlyknowndistributionofPassifloraaffinis inTexasisshowninFig..3,basedonherbariumandliteraturesurveys,indicat- ingthatitisnearlyrestrictedtothesouthernEdwardsPlateauandLampasas CutPlains(Diggsetal.1999;Turneretal.2003). ApopulationwasoncecollectedniFayetteCounty,Texas,byBennoMatthes (seespecimensexamined),representingtheeasternrange-limitofthespecies intheUnitedStatesandinanareawithdifferentgeologythanwherethisspe- ciesoccurselsewhereinTexas.Matthes,aGermansettlerandnaturalistin FayetteCounty,livedforashortperiodinthetownofRoundTop,wherehe collectedseveralplantspecimensfromlate1853untilmid-1854(Geiser1941). InthisareaheapparentlycollectedP.ajfinis(=Matthes274),withspecmiens depositedat the Museum National d^fiistoire Naturelle, Parisand the NaturhistorischesMuseWum,Vienna(Kiliip1938).ThissWpecimenstillexistsatP, butItwasnotfoundat inarecentsearch,although doeshaveafragmen- taryMatthesspecimenofPIutcalabeled"beiNo.274,"suggestingthatthetwo specieswereprobablygrowingtogetherN-iatthesinitiallyoutlinedhisbotani- caldiscoveriesinthisarea,althoughwithoutmentioningPassiflora(Matthes 1855a).Helaterprovidedamoredetailedfieldaccount(Matthes1861)wherehe mentionedfindingattheuppermarginofariparianforest'''Passifloratrilohd" with"Clematiscoccinea"andCornusjlorida.Thismayhavebeentothesouth- westofRoundTopnearCummins("Cummings")Creek,alongwhichhehad beenknowntocollect(Matthes1855b,1861).Althoughmesicforestedgesare appropriatehabitatforbothPaffinisandPlutcu,suchahabitatattheinner edgeofthecoastalplainmayseemmoreappropriateforthelatterspecies(e.g. Cooperrider1995;Alford2000)versusthoseofPaffinisontheEdwardsPla- teauandassociateduplift. However,muchofFayetteCounty,includingthe RoundToparea,isunderlain bysandstoneandclayof theCatahoulaand Oakvilleformations,bothwhichcancontaincalcareousportions,particularly thelatter(Sellardsetal.1932;Proctoretal.1974).Furthermore,thespringsin FayetteCountyarealsoknowntobealkaline(Brune1981).Thissuggeststhat muchofFayetteCountycanprovidehabitatsconsistentwiththeprimarily calciphilicprelerencesofPaffinis. 280 BRIT.ORG/SIDA21(1) 105° 100° 95° Fig.3.GeographicdistributionofPassifloraaffinisintlieUnitedStatesbasedontierbariumsurveysandfloristicsum- maries(Diggsetal.1999;Turneretal.2003). Killip(1938)reportedthepresenceo'lP.affinismNewMexico,citingaParry specimenatUSfromDonaAnacounty,laloeled"cinieflyinthevalleyoftheRio Grande,belowDonana."Thisspecimenwascollectedbythebotanicalsurvey partyattachedtotheUnitedStatesandMexicanBoundarySurvey,consisting ofJohnBigelow,CharlesParry,ArthurSchottandCharlesWright(listedonthe collectionlabelas"C.C.lAn-ryJ.M.Bigelow,C.Wright,&A.Schott":seespeci- mensexamined).1lowcver,theonlyP.affiniscollectioncitedmthebotanical reportoftheBoundarySurveyisbyBigelowinOctober(butwithnoyeargiven) alongtheDevilsRiver(Torrey1859)mwhatisnowValVerdeCounty,Texas, perhaps350milesfromDonaAnaCounty,NewMexico.TheBoundarySurvey travelledwithincloseproximitytotheRioGrandeforseveralhundredmiles southandwestol DonaAnaCountyaftertheTreatyofGuadalupeHidalgo GOLDMAN,LECTOTYPIFKATIONOFPASSIFLORAAFFINIS 281 wassignedwithMexicoin1848,followingtheendoftheMexican-American War(Emory1857).Thereforeitshouldnothavebeenassumedthatthisspeci- menwascollectedwithinthepresentboundariesofNewMexico. Incontrast,Killip(1938)didnotsimilarlyinterpretthevaguelabelsofP affinisspecimenscollectedbyCharlesWrighttosuggestthatthisspeciesoc- curredinNewMexico.Threecollectionscouldhavebeeninterpretedassuch, oneatGH(Wright217)labeled"fromWesternTexastoElPaso,NewMexico, MaytoOctober1849,"anotheratGHandUSwiththesameinformationbut numbered218,andanotherwithnonumberatNYandPHlabeled"N.Mexico, 1851"(seespecimensexamined).Itissurprisingthattheywerenot,withElPaso attributedtoNewMexicointwoofthesethreecollectionsand"N.Mexico"also potentiallyinterpretedmthesameway,oras"northernMexico." Wright'snumber217isactuallyadistributionnumbergivenbyAsaGray forhisPlantaeWrightianaepartI(1852),whereasWright'sfieldnumbercorre- spondingto217was40'1-,collectedonthe9^'"^ofJune,1849. Histravelroute (Geiser1935)andhisfieldnotes,summarizedinShaw(1987)andavailablein theGrayHerbariumarchivesatHarvardUniversity,indicatethat217/'104was collected"onthebanksoftheLeona"closetopresent-daytownofUvalde,Texas. Wright218wasthedistributionnumberforWright'sHeldnumber734,col- lectedonthe25*^^ofJuly1849,amongthe"hillsoftheDevil'sRiver"inpresent- dayValVerdeCounty,Texas,althoughGray(1852)consistentlyreferredtothe DevilsRiverasthe"SanPedroRiver",followingthegeographicnamesusedby French(1850;Shaw1987).Wright's"N.Mexico,1851"maybethesameasthat referredtobyGray(1853),collected"ontheSabinal,westernTexas,July,"inwhat isnowUvaldeCounty.However,Gray(1853)givesnodistributionnumberfor thisspecimen.InWright's1851fieldnotesintheGHarchivesPassifloraisen- teredonlytwice,oncehavingbeenfoundonChiconCreek(inmodernZavala County,Texas)inMay1851andanothertimefoundmthe"mountainsnearSanta Cruz,climbingamongrocks,"inSonora,Mexico,July1851,withnomentionof thegenusinhis1852notes.TheChiconCreekPassifloramatchestheentryin Gray(1853)forP.tenuilobaandtheSantaCruzspecimenmatchestheentry (andprotologue)forP.inamoena(=P.hryonioidesKunih).InJuly1851Wright washundredsofmilesawayfromtheSabinalRiver,inthevicinityof ElPaso andsouthwesternNewMexico(Shaw1987).Itispossiblethathecollectedthis Paffinisspecimenmthisarea,butwithoutenteringitintohisfieldbook,al- thoughthatseemsunlikelybecausehewasafastidiousnote-takerduringhis explorationsoftheborderregion.Theonlytimeheappearedtorelaxhisnote- takingwasattheendofhistravelsin1852,onhisreturntoSanAntonio,when onJuly2'^'-^herecordshisfinalcollectioninwhatisnowwesternCrockett County,Texas(Shaw1987),over100milestothenorthwestottheSabinalRiver inUvaldeCounty.ItseemsprobablethatWrightcollectedthisspecimennotin 1851butin1852,eastofCrockettCountyonhisreturntoSanAntonio,which 282 BRIT.ORG/SIDA21(1) wouldhavetakenhimthroughpresent-dayUvaldeCounty.Shaw(1987)men- tionsthatthesepost-July2,1852notestendtobefolded-upinthefragment foldersofthespecimensthemselves.SuchnotestendtobeonlywithGHspeci- mensbecauseol Wright'sprofessionalassociationwithGray.Noduplicateof thisspecimenexistsatGHsosuchinformationisprobablylost.Furthermore, Grayoftencutupnotesandcorrespondencetomakeintofragmentfolders(W. Kittredge,pers.comm.).ThusifthesenotesstillexistatGHtheymayforma packetononeoftheoverfivemillionspecimensmtheherbarium. ThesethreeWrightcollectionsandthesingleParrycollectionwereob- tainedwithinwhatiscurrentlywest-centralTexas,someofthisareahaving beenincludedinasoutheastwardextensionottheMexicanprovinceofNew Mexicopriortothesigningof theTreatyot GuadalupeHidalgo(Commons 1990).Thereafter,allthatispresentlypartofmodernTexaswasconsideredto bewithintheUnitedStates,Texashavingcfaimedmostotwhatisnowmodern NewMexico,northwardthroughpartsolColoradoandKansas,intoWyoming, thislandlaterpurchasedfromTexasbytheUnitedStatesintheCompromise of1850(Stephens&Holmes1988;Beck&Haase1989).WiththeseParryand WrightcollectionsmadeafterGuadafupeHidalgoinanareathathassubse- cfuentlybeenconsideredtobepartofTexasonly,anyinterpretationsinher- bariumlabelsorintheliteratureofthesecollectionshavingcomefromNew^ MexicoornorthernMexicoareinerror. Despitethepotentiallymisleadinglabelsonthesespecimens,Raffinishas neverbeenreportedfrommodernNewMexicoinanyflorasofthatstate,which isfortuitousbecauseasimilarsituationwithvagueherbariumlabelsmayhave ledtothepotentialmisreportingofP.tcnuilobaforNewMexico(Martin& Hutchms1981).Therefore,itcanbeassumedthatthegeographicrangeol P. dffinisisrestrictedonlytocentralTexas,withscatteredpopulationsinnorth- easternMexico.Thepotentialconfusioncreatedl^yold,vaguelabels,canlead tofloristicuncertaintyandthisaloneshouldbeaninspirationtoallbotanists tobeextremelythoroughinmakingtheircollectionlabels. Specimense.xami[ied:MEXICt).NuevoLeon.Mcpo.LanipazasdeNaranjo:RanchoResendez, Lampazas,22Jun14^7.M7.]:(lw(Uils29S(GH).Mcpo.Monterrey:MoniciTC)'.SierraMadreC^ricnial, 23Jul1933,(IHMucUc)&Ml.MuellerJOOCGHlMcpo.Santiago:trailbetweenPotreroRedondo& LC:aesrrAojuZnatmaosr,a2,422AuAgug1Q1390,30C..Ht.ilMluBUaretrlct2t95H509(G4HJ(.GHT)a.mMacuploi.paVsicMtocrpioa.:IC2:irnuiillWas:ofVicV.icotforEila,M2i8lagAruog, 1948.Ki-Mdver&Crum.5366(GtU, U.S.A.TEX/\S:BexarCo.:ComancheSpring,NewBraunlels,etc.,Aug-Sep,1849,h]Lindbeimer 817N74I(ARIZ.BM,BR.BR1T-SN4U,C,CAN,G[2lGH[21K,MO.NY,RPH,TEX,DC.US,W);San Antonio,17Oct1920,E.D.Sthide401(LISJ.EdwardsCo.:DevilsSinkHole.29Sep1940,V.LCoyy X5652(GH);39miS,ofSonora,l(iAug1941,V'L,Cory,58039(G\\):DevilsSinkHole,lOmiH,olRock Springs,24Jul1946,DS.CondiIM12(BRIT,NY);lOmiSEofCartaValley,7Aug1965.D.S.Cortxll 31.537(GH),FayetteCo.:nolocalit\,nodate,RMcfiflies274(P),GillespieCo.:3iTnE,offarmroad.Imi S,ofcountylineoffn.lb,20Jul1967,DS.Correll&H.B.ConcU,H292iGlI).HaysCo.:SanMarcos GOLDMAN,LECTOTYPIFICATIONOFPASSIFLORAAFFINIS 283 andvicinity.Jul1897,S.W.Stanfiehls.n.(NY):lakebank,SanMarcos,25Jul1941,B.C.Tharps.n.(BRIT, GH,NY,PH,USj.KerrCo.:Hunt,shoresofGuadalupeRiver,4Aug1969,P.Fryxell1099(NY).Kimble 3Co1.5:2T7e(lGeHg)r.apKhi,n8neOyctCo1.91:6F,t.E.jC.lParakl,m1e0rM10a9y4218(9U3S,);E.aAl.oMnegaLrlnasno14R3i9ve(rUSa)t;Ju1n0c3ti/o4nm,i6WSepof19L6a3g.uDn.aS.,C2o4rSredpI 1939,V.LCory33461(GH).LlanoCo.:EnchantedMt.,Fredricksburg,26Jun1932,CC.A/Ixt.s.s.n.(BRIT); EnchantedRock.1Aug1938,B.C.Tharps.n.(NY).SuttonCo.:3/4miSWofFt.Terrett,20Aug1937, V.L.Cory24090(GH).TravisCo.:aboveZilkerPark,Austin,10Oct1945,B.C.Tharp45-49(GH,NY); HamiltonPool,8Jul1966,J.R.Crulchjield1802(NY):Austin,27Aug1978J.M.MacDougal448(US). UvaldeCo.:fromWesternTexastoFlPaso.NewMexico.May-Oct1849,CWright217(GH|2]);N. Mexico,1851,C.Wrights.n.(NY,PH).ValVerdeCo.:chieflyinthevalleyoftheRioGrande,below Dofiana,nodate,C.C.Parry.J.M.Bigelow.C.Wright.&A.Schotts.n.(US):fromWesternTexastoEl Paso,NewMexico,May-Oct1849,CWright2J8(GH[21US),Undeterminedlocality:Nueces,1880s,V. Havard.s.n.(US).CULTIVATED:CornellUniversity,originallyfromU.Texas.SanAntonio.17Jun2001, D.Goldman1769(BH);CornellUniversity,originallyfromU.Texas,SanAntonio,10Aug2001.D. Goldman1819(BH);CornellUniversity,originallyfromU.Texas,SanAntonio,9Sep2001,D.Gold- man2126(BH);CornellUniversity,originallyfromEdwardsCo.,TX,9Sep2001.D.Goldman2127 (BH);CornellUniversity,originallyfromU.Texas,SanAntonio,28Sep200f,D.Goldman2160(BH); CornellUniversity,originallyfromU.Texas,SanAntonio,2Nov2001,D.Goldman2161(BH);Cornell University,originallyfromU.Texas,SanAntonio,2Nov2001,D.Goldman2162(BH). ACKNOWLEDGMENTS Th£\nksisgiventoLisaDeCesare,GretchenWadeandJudyWarnementforli- braryandarchivesassistance;KenCameron,KanchiGandhi,WakerKittredge, Scott"M"LaGreca,BarneyLipscombandTanyaLivshultzforvaluablediscus- sionsorassistance;JohnMacDougalandKristenPorter-Utleyforreviewingthis manuscriptandotherrelateddiscussions;thestaffatARIZ,BM,BR,BRIT-SMU, C,CAN,G,GH,K,MO,NY,RPH,TEX,UCandUSforsendingloansand/or makingdigitalimagesofspecimensavailable;Alain Changy(P),Jochen Heinrichs(GOET)andBrunoWallnofer(W)forassistancewithfindingMatthes specimens;BobDirig,KevinNixonandSherryVance(BH),andRichardZander (BUF)forreceivingloansonmybehalf;MarshallEnquistandLarryGilbert (TEX)forsendinglivematerialforcultivation;CarolBader,PaulCooper,Jerry David,KimGoodwin,AndyLeedandMelissaLucknowforthefacilitationof spaceandcultivationassistanceatCornellUniversity;AndrewColliganforas- sistancewiththearchivesoftheMissouriBotanicalGarden;RenateHellmiss andDanielSchottfortranslationsofMatthesmanuscripts;DanFulopforthe SpanishtranslationoftheabstractandHobbesGoldmanforgeneralsupport throughout. 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