OSMORHIZABIPATRIATA(APIACEAE)INTEXAS: TAXONOMICSTATUSAND CONSERVATIONCONSIDERATIONS JunWen PorterRLowryII DepartmentofBotany MissouriBotanicalGarden TheFieldMuseumofNaturalHistory P.O.Box299 1400S.LakeShoreDr. St.Louis,Missouri63166-0299,U.S.A. Chicago,Illinois60605-2496,U.S.A and and DeparterventdeSystematiqueetEvolution LaboratoryofSystematic&Evolutionary MuseumNationald'HistoireNaturelle Botany&Herbanum,InstituteofBotany CasePostale39,57rueCuvier ChineseAcademyofSciences 16rueBuffon Beijing100093,CHINA;[email protected] 75231Paris,FRANCE;[email protected] JamesC.Zech DepartmentofBiologyC-57 SulRossStateUniversity Alpine,Texas79832,U.S.A.;[email protected] ABSTRACT ThepaperevaluatesthetaxonomicandconservationstatusofOsmovhxzaUpatriataConstance& R.H.Shan.Theavailablephylogeneticdatasupportitsrecognitionasadistinctspecies,ithasahighly restricteddistributiononMt.LivermorcoftheTransT'ecosregionmTexas,aswellasinCoahuila andNuevoLeon,Mexico.ItshabitatonNit.LivermoreishighlyrestrictedwithinthemesicPifius strohijormiscommunitywhereitisassociatedwithseveralothertaxathatarealsorareinTexas, suchasPohmoniur}!paucijlorumS.Watsonsubsp.bmcklcyi(Standi.)WherryandAquilev,ia longissimaA.GrayThepopulationofO,hipalnalcionMt.Livermoreissmall(<20individuals), althoughapparentlyhealthy,withafullrangeofagestructure.ApplicationofthelUCNRedList criteriashowsthat0.hipatriataisanendangeredspecies[B2ab(iv)].Werecommendcarefulmoni- toringofO.hipalriataanditsassociatedhabitat,andrestrictedaccesstoitsonlyknownlocality withintheUnitedStates. RESUMEN EnestearticuloseevaluaelestatustaxonomicoydeconservaciondeOsmorhizabipairiataCon- stance&R.H.Shan.Losdatosfilogeneticosdisponiblesapoyansureconocimientocomounaespecie independiente.TieneunadistribucionrnuyresiringidaenelMt.LivermoredelaregionTrans-Pecos enTexas,aslcomoenCoahuilayNuevoLeon,Mexico.SuhabitatenelMt.Livermoreestarnuy restringidodentrodelacomunidadmesicadePinusslrohiformis,dondeestaasociadoconotrosvaries taxaquetambiensonrarosenTexas,tale;comoFolemoniurnjid11ci/lori(mS.Watsonsubsp.Iiinc/dcvi (Standi.)WherryyAquilegialongnsimaA.GrayLapoblaciondeO.bipcitriataenelMt.Livermorees pequefia(<20individuos),aunqueaparentementesaludable,conunaestructuradeedadconelrango complete.LaaplicaciondeloscriterioslUCNparalaListaRojamuestranqueO.hipatnaiaesuna especieamenazada[B2ab(ivj].Recomendamosuncontrolcuidadosode0.hipatriataysuhabitat asociado.yaccesorestringidoasuunicalocalidadconocidaenlosEstadosUnidos. SIDA21(1):501-506.2004 502 BRIT.ORG/SIDA21(1) INTRODUCTION OsmorhizaRaf.(Apiaceaesubfam.Apioideae)isasmallgenusofapproximately 11species,includingninethatoccurinNorthAmerica(twoofwhicharealso disjunctinSouthAmerica)andoneeachmAsiaandthecentralAndes.Mem- bersofthegenusarecharacterizedbytheirdeepbrowntoblack,lineartoob- long,fusiformtoclavatefruitsthatareslightlycompressedlaterallyandhavea caudatebasalappendageandfiliformribs,whichinmostspeciesbearnumer- ousretrorsebristles(Lowry&Jones1984).Severalrecentmolecularphyloge- neticanalyseshaveshownthatOsmorhizaismonophyleticandthatitismost closelyrelatedtotheOldWorldgeneraMyrrhisMill,andGcocaryumCoss. (Downieetal.2000;Wenetal.2002;Yooetal.2002).Thesestudieshavealso indicatedthatalltheNewWorldmembersofOsmorhizaevolvedfromacom- monancestorthatarrivedfromAsia,wherethegenusappearstohaveorigi- nated(Wenetal.2002).WithintheNewWoddclade,agroupoflargelywest- ernNorthAmericanspeciesformsawellsupportedsubcladethatprobably diversified relativelyrapidlyTwoofthesespecies,0.herteroi DC.and0. depauperataPhil,havesubsequentlydispersedintotheGreatLakesarea,east- ernNorthAmerica,andsouthernSouthAmerica,andanother,0.mexuana Griseb.,hasaninterruptedrangefromnorthernMexicotonorthernArgentina, whiletheremainingspeciesoccupyrelativelynarrowgeographicrangesinthe westernUnitedStates(Lowry&Jones1984;Wenetal.2002). Specimensattributedtooneofthesespecies,OsmorhizabipatriataCon- stance&R.H.Shan,werefirstcollectedonSeptember28,1935,byL.C.Hinckley onMt.LivermoreintheDavisMountainsoftheTrans-PecosregioninTexas (Hinckley408,SRSC).Constance&Shan(1948)described thissweetcicely (hereinreferredtoasthebipatriatesweetcicely)fromtheDavisMountainsand northernMexicoasaspeciesnewtosciencebasedonalatercollectionby Hinckley(26Jul1937,LC.Hinckleys.n.;holotypeatNYandisotypesatARIZ andGH).Constance&Shanchosetheepithettoreflectthepresenceof0. &bipJaotnreisata(1m98t4w)orceopuonrttreidessaesvewrealllamsotrhpehotlwoogniactaiolnailnitteiersmeofdiiatstaeusthboerts.weLeonwr0y. bipatriataandO.mexicanacollectedfromsitesinnorthernMexico,including onCerroPotosiinNuevoLeon,wherethetwotaxaoccursympatricallyThese intermediatespromptedthem toreduce0.bipatriata toasubspeciesofO. mexicana.RecentlyTurnerctal.(2003)recognizedthetaxonasavarietyof0. mexicanabecausehechose,somewJiatarbitrarilynottousetherankofsub- speciesinhiswork(B.L.Turner,pers.comm.). TAXONOMICSTATUS Phylogeneticanalysesconductedmthelastseveralyearsusingsequencedata fromthenuclearribosomalITSregionsandthechloroplastnx/hFgeneandtrnL- WENETAL.,OSMORHIZABIPATRIATAINTEXAS 503 Fregions(Downieetal.2000;Wenetal.2002;Yooetal.2002)suggestthat Osmorhizcimexicanaand0.bipatriatadonotformamonophyleticgroupand indicatethattheyshowconsiderabledivergenceintheirsequenceprofilesof thesemarkers(e.g.,1.415%divergencebetweentheITSsequences).IntheITS phylogeny,thepositionsofthetwotaxaareunresolved,althoughbothare closelyrelatedtootherwesternNorth AmericanmembersofOsmorhiza (Dowmeetal.2000;Wenetal.2002).Thechloroplastdatasetsuggeststhat0. hipatriataismostcloselyalliedtoO.depauperata(Yooetal.2002).Theavail- abledatathusdonotsupporttheinclusionof0.hipatriatawithinO.mexicana ataninfraspecificlevel.Thephylogeneticdatasuggestinsteadthatitshould betreatedasadistinctspecies,asinitiallyproposedbyConstance&rShan(1948), usingeitherthephylogenetic(Nixon&Wheeler1990)orthelineage(deQueiroz 2000)speciesconcept.Whilethepresenceofmorphologicalintermediatesin northernMexicowouldappeartobeincompatiblewiththerecognitionof0. hipatriatausingtheclassicalmorphologicalspeciesconcept,assupportedby Cronquist(1978),thephylogeneticdatasuggestthattheseintermediatesmay representinterspecifichybridsorperhapsmayhaveresultedformintrogres- sionbetween0.hipatriataand0.mexicana.Detailedanalyses,however,will berequiredtoassesstheoriginoftheseintermediates. Osmorhizahipatriatadiffersfromitscongenersbyitssmall(9-11mmlong) andglabrousfruits,whichareImear-fusiformandtaperintoashortbeakat theapex,andbyitsveryshortstyles[0.5-0.75mmlong(Lowry&Jones1984)1 Basedprimarilyonfruitmo.'phology,Constance&Shan(1948)placed0. hipatriatainsubgen.Glycosma,butcommentedthatitpossessedcharacters intermediatewithmembersofthetypicalsubgenus,inparticularwithregard toitsinvolucel,stylesandstylopodium,whicharesimilartothoseofO. mexicana,whichbelongstosect.Mexicanaeofsubgen.Osmorhiza(seeCon- stance&Shan1948;Lowry&Jones1984).Ourmolecularphylogeneticstudies (Wenetal.2002;Yooetal.2002)showthatO.hipatriatapossessesalargenum- berofuniquemolecularnucleotidesubstitutions,suggestingitsantiquity. Osmorhizahipatriataisthusperhapsbestdescribedasarelictspecies,which hassurvivedinmesichabitatsInthenortheasternChihuahuanDesert,which haveundergoneagradualdryingsincethelateTertiary(Graham1993). AnillustrationofOsmorhizahipatriataisfoundinLundell&collaborators (1961:plate42).Lowry&Jones(1984)providedcomprehensiveinformationon itsmorphologyanddistributionandcitedspecimensfrombothTexasandMexico depositedinvariousherbaria.BelowaresomeadditionalspecimensfromTexas thatwehaveexamined,butwhichwerenotseenbyLowry&Jones(1984). U.S.A.Texas.JeffDavisCo.:MaderaCaryon.Ml.laverrnore,28Sep1935.infr..Hintkhy408iSRSC)\ inwetsoilofspringinMaderaCanyon,Mt.Livermore,23Jul1945.inyoungfr,Hinckley3489iSRSC); attheupperspringofMaderaCanyon,NWslopeotMt.Livermore.7300ft,12Sep1947,inIr.,Warnock &Hinckley4M7[SRSO:rareatupperspringinMaderaCanyononMt.Livermore.DavisMountains, 504 BRIT.ORG/SIDA21(1) igneoussoil,7500ft,11Sep1947,infr.Warnock&Hinckley7479(SRSC);infrequentinigneoussoil eornauCpapneyronM,adTeorbaeCSparniynog,nNofWMto.fLMitv.eLrimvoreer,mo7r3e0.0jufts.t2a3boA\'uega1lQa6r8g,eWhaorrsneoscfkio2e3w0aJl5l,(SinRSdCe)e;pumpepseircMcaadn-- yonunderPmusstivhifoimis-Qucicus^amhcUi-jumpciusdcppcanalorest,associatedwithRham- nushetulijolia,Symphoricarposoreophilus,Aquilegialongissima.Polcmoniumpaucijlorumsubsp. hinckleyi,Salviaarizonica.Geraniumcaespitosa.Vitisarizonica.Pteleatrifoliata.Galiumsp..and Thalicirumsp.,N30°38.364',VV104°10.746',7242ft.,23Aug2003,\nfl&fr,Wenelal.7265{P}. CON.SERVATIONC0NSIDERAT10N,S Osmorhizahipatriatahasahighlyrestricteddistribution,withonlyafewlo- calitiesrecordedonMt.Livermore,Te.xas,andinCoahuilaandNuevoLeon, Mexico.Untilrecently,allofthecollectionsIromTexasweremadebyL.C. HinckleyandBartonWarnock(perhapsfromthesamelocality),themostre- centofwhichdatesfrom1Q68(Warnock23015,SRSC).Duringarecentstudy trip,wewereabletorelocate0.bipatricitaonthenorthwesternslopeofMt. Livermore.Asingle,smallpopulationwasfoundon23August2003,compris- ingabout15individuals,including5-6youngseedlings,itoccupiedasmall microhabitat withmoist,rich soil in adeepmesiccanyonunderPmus strohiJormis-Quercusgamhelii-Juniperusdeppeanaforestthatdiffersconsid- erablyfrommostofthesurroundingvegetation,whichisprimarilydominated byponderosapinesandvariousoakspecies.TheplantsofO.hipatriataonMt. Livermorewerehighlyheterogeneousphcnologically,withsomeindividuals observedinflower,someinyoungfruit,andothersinmaturefruit,inaddition toalewthatwerejustemergingfromthegroundafterarecentrainshower, whichsuggeststhat0.bipatricitaisparticularlysensitivetowateravailability. SuitablehabitatforOsmorhizahipatriataisapparentlyrareintheDavis Mountains.Althoughthepopulationwestudiedappearstobehealthy,asindi- catedbythepresenceolafullagestructure,fromyoungseedlingstomature adultplants,itissmall(lessthan20individuals)andonlypersistsinarestricted microhabitat.Despitethreedayssearchingthemountainforcomparablehabi- tat,wewereunabletolocateanyotherpopulations.Wethusrecommendcare- fulmonitoringoftheonlyknownpopulationintheUnitedStates,whichmay requirecontrolledaccesstoensureitscontinuedsurvival.Osmorhizahipatriata alsooccurssympatricallywithtwootherraretaxaintheDavisMountains:Po- kmoniumpaucijlorumS.Watsonsubsp.hinckleyi(Standi.)Wherry;andAq- uilegialongissimaA.Gray.Wealsoobservedtwoadditionalrarespecies(Qucrcu.s depressipesTrel.andAraliahicrenataWooton&Standi.)atahigherelevation onthesameslopeofMt.Livermore,thesummitoftheDavisMountains. OsmorhizahipatriataisnotcurrentlylistedintheTexasThreatenedand EndangeredSpecieslist(TexasParks&WildlifeDepartment2003)norinthe ThreatenedandEndangeredSpeciesSystem[TESS(U.S.FishandWildlife Service,2003,http://endangered.fws.gov)].ApplicationofthelUCNRedList criteria(lUCN,2001;seealso http://www.iucn.org/themes/ssc/redlists/ WENETAL.,OSMORHIZABIPATRIATAINTEXAS 505 RLcats2001hooklet.html)shows,however,thatO.hipatriataisanendangered species[B2ab(iv)]becauseithasanareaofoccupancyestimatedtobelessthan &50J0oknmes^1a9n8d4:beficga.u1s7e)iatnisdbaoptphekanrsowtonhfarvoemafedweecrhtnhianngfnivuemlboecarhtoifessu(cbfp.oLpouwlray- tions,asindicatedbyourobservationsonMt.Livermore. MostspeciesofOsmorhizahavearelativelywidedistribution(cf.Lowry &Jones1984).AlongwithO.glahrataPhil.,acentralAndeanendemic,andO. hrachypodaTorn,foundonlyinCaliforniaandArizona,O.hipatriatahasone ofthemostrestrictedrangesinthegenus.Inthecaseof0.hipatriata,thismay atleastinpartreflectthefactthatitsfruitslackthecaudateappendagesand retrorsebristlescharacteristicofmostmembersofthegenus,whicharepre- sumedtoplayaroleindispersal(Constance&rShan1948;Lowry&Jones1984), assuggestedforthewidespread0.herteroi,whichCruden(1966)concluded wasbirddispersed. Inconclusion,Osmorhizahipatriataisadistinctspecieswithalongevo- lutionaryhistory.ItappearstobeamongtherarestplantspeciesintheUnited States,withasingleknownpopulationoflessthan20individualsoccurringin ararehabitatintheDavisMountainsofwestTexas.Carefulstudiesneedtobe conductedtoinvestigateitsreproductivebiologyseedgerminationanddis- persalbiology,ecologicalpreferences,andconservationgeneticstodevelopa suitablemanagementplanforthisrelictspecies.Comparativeanalysesofits congenerswithbothwidespreadandrestricteddistributionsshouldalsopro- videinsightsintothebiologyof0.hipatriatathatmaybeusefulforitsconser- vation.Furthermore,itisnecessarytocomparetheTexaspopulationwiththose inCoahuilaandNuevoLeon,M^exicoinordertoverifythattheyareindeedcon- specific,assuggestedbytheirmorphology.Theevolutionaryconsequenceor impactofthepossiblehybridizationandintrogressionbetweenOsmorhiza hipatriataandO.mexicanaalsoneedstobeassessed. ACKNOWLEDGMENTS WethanktheNatureConservancyofWestTexasforgrantingaccesstotheDavis MountainsPreserve,authorizingourfieldstudyonMt.LivermoreinAugust, 2003,andprovidinglodgingduringthefieldwork;wearegratefultoJohnKarges forhelpfuladvice,PatriciaManningforfieldassistance,MikePowellforher- bariumassistance,permissiontoexaminethespecimensatSRSC,andvaluable commentson the manuscript,B.L.Turnerfordiscussionand hisopen- mindednessofacceptingthespeciesstatusofOsmorhizahipatriata,andGeorge SchatzforassistancewiththelUCNRedListanalysis.Supportforthestudy wasprovidedbytheFieldMuseumofNaturalHistory,theChineseAcademy ofScience(agrantofExcellentOverseasChinese),andtheNationalScience Foundation(DEB0108536toJV/). . 506 BRIT.ORG/SIDA21(1) REFERENCHS Constance,L.and R.H.Shan.1948.ThegenusOsmorhiza(Umbeliiferae),astudyingeo- graphicaffinities,Univ.Calif.Publ,Bot.23:ln-156. Cronquist,a.1978.Onceagain,whatisaspecies?In.-L.V.Knutson,ed.Biosystematicsin agriculture.AllenheldOsmun,Montclair.Pp.3-20. 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