ENDEMICVASCULARPLANTSOFTHEINTERIOR HIGHLANDS,U.S.A. DouglasZollner MichaelH.MacRoberts TheNatureConservancy BogResearch 601NorthUniversity 740Columbia LittleRock.Arl<ansas72205,USA. Shreveport,Louisiana71104,U.S.A. Barbara R.MacRoberts DouglasLadd Herbarium,MuseumofLifeSciences TheNatureConservancy LouisianaStateUniv-Shreveport 2800S.Brentwood Shreveport,Louisiana71115,U.S.A. St.Louis,Missouri63144,U.S.A. ABSTRACT Weevaluatethe36cndemievascularplatitsoftheInteriorHighlancisolArkansas,Illinois,Kansas, Missouri,andOklahoma.Mostoftheendemicfloraoftheregionareherbaceousperennials,although nearlyaquarteroftheendemicplanttaxaareannuals.AnanalysisofthecommunityaiIiliationsof theregion'sendemic1lorarevealsthatmostendemictaxaareassociatedwithgladehabitatsmain- tainedb)'edaphicconditionsandfire.Riparianhabitatsandlire-maintained]iine-oakwoodlands areotherliabitatassociationswithsignilicantendemicIlora.TheOuachitaMountainsregionolthe InteriorHighlandscontainsadisproportionaterepresentationoltheendemictaxarelativetoitsarea, althoughthereisalsoasmallercohortofendemicsrestrictedtotheOzarkportionoltheInterior Highlands. Khywords:InteriorIlighlands.endemicIlora,biogeography,biodiversit)',Ozarks,Ouachitas,Arkan- sas,Illinois,Kansas,Missouri,Oklahoma RESUMEN Hemosevaluadolas36plantasvascularesendemicasdelasInteriorIlighlandsdeArkansas,Illinois, Kansas,Missouri,yOklahoma.Lamayorpartedelafloraendemicadelaregionsonherbaceasperennes, auncjuccasilacuartapartedclasplantasendemicassontaxaanuales.Unanalisisdelasprelerencias delascomunidadesdelailoraendemicadelaregi6nrevelaquelamayoriadelosta.xaendcmicosestan asociadosconhabitatspantanososquesemantienenporlascondicionesedaiicasyeltuego,Los habitatsripariosybosc|uesde|iino-roblemantenidosporelfuegosonlosotroshabitatsconelementos defloraendemicasignificativos.LaregiondelasMontanasdeOuachitadelInteriorHighlands contieneunarepresentaciondesproporcionadadetaxaendemicosdeestaarea,aunquehaytambien unapequenacohortedeendemismosrestringidosalapartedeOzarkdelasInteriorHighlands. INTRODUCTION Listsofendemictaxaarevaluabletoolsfordefiningtheuniquenessandcon- servationsignificanceofbiogeographicregions(Takhtajan1986)andinestab- lishingbiodiversity"hotspots"forconservationpriorities(Diamondetal.1997; Rickettsetal.1999;Myersetal,2000;Stemetal.2000;Estill&Cruzan2001). Fewbiogeographicregions,however,havedetailedendemicvascularplantlists. SIDA21(3):1781-179L2005 1782 BRIT.ORG/SIDA21(3) Forexample,onlyrecentlyhaveendemicplantlistsbeendevelopedfortheAt- lanticandGulfCoastalPlainandtheWestGulfCoastalPlain(Sorrie&Weakley 2001;MacRobertsetal.2002). TheInteriorHighlandsphysiographicregionhasbeenrelativelywell-de- scribedphysicallyandbiotically(Fenneman1938;Foti1974;Nelson1985;Thorn &Wilson1980;Bryantetal.1993;NighandSchroeder2002;Skeenetal.1993; Foti&Bukenhofer1998;Delcourt&Delcourt2000).Theregionhasanexten- sivebotanicalliterature,includingIlorasandatlases(Steyermark1963;Peck& Peck1988;Smith1988,1994;Taylor«&rTaylor1989;Pecketal.2001;Yatskievych 1999),butithasnocomprehensivelistolendemicvascularplants,andmost botanicalinformationiscompartmentalizedbystate. AsshowninFigure1,theInteriorHighlandsarecomprisedoftheOzarks, Ouachitas,andArkansasValleysections,andincludesignificantportionsolAr- kansas,Missouri,andOklahoma,alongwithminorareasofIllinoisandKansas. Thisregionhaslongbeenrecognizedasageologically,physiographically,eco- logically,andculturallydistinctregionofNorthAmerica,andconstitutesthe onlyhighlandsinmidcontinentalNorthAmerica.TheOuachitaMountainsare east-westtrendingfold-beltrangesofintenselydeformedsandstone,shale,and chert(Miser1929).Thenorthernrangesarelonghogbackridgesotsandstone separatedbybroadvalleys.Thesouthernrangesaresharpridgesofnovaculite separatedbynarrow,stonyvalleys(Croncis1930).TheArkansasValleyisabroad alluvialplainwithisolatedmountainsthatgenerallyseparatestheOuachita MountainsfromtheOzarkPlateau.Structurally,theOzarkPlateauisadomethat hasbeenslowlyupliftedanderoded,resultinginhighlevelsoftopographic,geo- logic,edaphic,andhydrologicdiversity.Bedrockgeologyincludesexposuresof PrecambrianigneousrockssurroundedbyalternatingzonesofPaleozoicsand- stoneandcarbonatesedimentaryrocks(NighandSchroeder2002).Boththe OuachitaandOzarkportionsofthehighlandsarccharacterizedbyrugged,dis- secteduplandswithabundantexposedrocksandhighlyvariablesoildepths. ThispaperenumeratestheendemicvascularplantsoftheInteriorHigh- lands,alongwithmtormationabouteachspecies,itsplantcommunityaffilia- tionsandtheecologicalprocessesthatmaintainthesespecies.Thesedatacan beusedtofocusactivitiesonthehabitats,ecologicalsystems,andecological processregimesingreatestneedofconservationaction.Gladeisusedhereto mean,openherbaceous-dominatedhabitatswithsparsetreeandshrubcover, shallowsoils,andabundantexposedrock. MHTHODS Forthisstudy,ataxonisconsideredtobeendemicifitsrangeessentiallydoes notextendoutsidetheInteriorHighlands.Hybridtaxaareexcludedfromthis list.NomenclaturegenerallyfollowsKartesz(1999). Wesearchedallavailablesourcesofinformation,includingextensivecon- ZOLLNERETAL,ENDEMICPLANTSOFTHEINTERIORHIGHLANDS 1783 sultationwithknowledgeableexperts,todetermineglobalrangesofspeciesin thevascularfloraofArkansas,Illinois,Kansas,Missouri,andOklahoma.This includedgeneralreferencessuchasKartesz(1999),Krai(1983),Mohlenbrock (2002),RobisonandAllen(1995),Smith(1988),Steyermark(1963),Taylorand Taylor(1989),Yatskievych(1999),FloraofNorthAmericaEditorialCommittee (1993-),andmorespecificpapers(e.g.Tucker1974;Weckman2002).Alsoin- cludedwerevariouslistsofspeciesofconcernkeptbytheArkansas,Illinois, Kansas,Missouri,andOklahomaNaturalHeritageprogramsandtheOzark, Ouachita,MarkTwain,andShawneeNationalForests^. Oncethepreliminarylistwascompiled,weinvestigatedmanyofthesame sourcesandadditionalecologicalreferencestodeterminetheplantcommunity affiliation(s)ofeachspeciesthroughouttheInteriorHighlands.Thisdetermi- nationinvolveddevelopingageneralplantcommunitylistfortheInteriorHigh- landsandassigningeachtaxontothemostappropriateplantcommunity.We usedTheNatureConservancy'sclassificationattheecologicalsystemlevel (Comneretal.2003),makingecologicalsystemconceptuallyanalogoustoplant communityasappliedhere.Sometaxawerecharacteristicallyaffiliatedwith twoormoreecologicalsystems.EcologicalsystemsoftheInteriorHighlands arelistedinTable2. RESULTS Includedbelow(Table1)isanannotatedlistofthe36vasculartaxaendemicto theInteriorHighlands,arrangedalphabeticallybyfamilyandgenus.Foreach taxon,oneormorecharacteristicplantcommunitytypesareindicatedinbrack- etsaftertheplantname,usingthenumbersdesignatedforeachcommunityin Table2.Foreachspeciesinthislist,wealsoindicatethefamily,physiognomic profile,distributionpatternwithintheInteriorHighlands-Ozarks(Oz), Ouachitas(Ou),orboth-andadditionalrelevantinformationandcomments whereapplicable.TheArkansasValleyisincludedintheOuachitasection. DISCUSSION Weidentifiedatotalof36endemicvasculartaxawithintheInteriorHighlands; therearenoendemicfamiliesorgenera.Thereare24endemicspecies,and12 'inadditiontothetaxatreatedinthispaper,sixspeciesofCrataegus(Rosaceae),CcamllensisSarg.,C. harveyanaSatg.,C.lanuginosaSafg.datebrosaSarg.,Ciuc/oSarg.,andCAhermopegaeaPalmer,arerecognizedby Kartesz(1999)asbeingendemictotheregionandotherpotentiallyendemichawthornshavebeenproposed {e.g.C^ouachitensisPalmer)butSmith(1994)doesnotrecognizethem.EventhoughendemismishighinCrataegus, wehaveleftthiscomplexandincompletelyunderstoodgenusoffthelist.TherearereportsofHamamelisvernalis Sarg.(Hammamelidaceae)fromsoutheasternTexasbyCorrellandJohnston(1970),Vines(1977).Nixon(1985), Joneset.al.,(1997),andTurneretal.(2003),althoughFloraofNorthAmerica(1997)statesthatthisspeciesis endemictotheInteriorHighlands.Consequently,wehaveomittedHamamelisvernalisfromthistreatment.A revisionofthegenusTallnum(Portulacaceae)currentlyunderwaymayresultintheadditionoftwoendemic speciestotheInteriorHighlandsflora. 1784 BRIT.ORG/SIDA21(3) FiG.I.MapoftheInteriorHighlandsPhysiographicRegion. endemicsubspeciesorvarietiesofmorewidelydistributedspecies.Mostofthe endemicplanttaxaintheInteriorHighlandsareontheArkansas,llhnois,Kansas, Missouri,and/orOklahomaNaturalHeritageprograms"speciesofconcern"lists. Most158%)of theendemicvascularplantsoftheInteriorhighlandsare perennialforbs;thisparallelsthephysiognomyoltheoverallvascularfloraof theregion,aswellastheccoregionsol midcontinentalNorthAmerica.Only threeendemictaxa(,8%)arcwoody,andthereisasingleendemiclern.Although onlynineendemictaxa[15%)areannuals,thisisahigherproportionofannu- alsthaninthetotalnativevascularfloraol theregion,andislikelyreflective oftheharshconditionsthatcharacterizegladeenvironments,whichharbora disproportionatecomponentol annualspeciesascomparedtootherhabitats intheregion. ZOLLNERETAL.,ENDEMICPLANTSOFTHEINTERIORHIGHLANDS 1785 Table1.EndemicvascularfloraoftheIrTteriorhighlandswithcommunityassociationsinbrackets (seeTable2),0u=OuachitaMountains,Oz=OzarkPlateau,both=bothsections. APOCYNACEAE CYPERACEAE Amsofiiahubrichtii\Noods.[]2]both;perennial Corex/afebracteafaWaterfall[5,6,12]Ou;peren- forb nialsedge ASTERACEAE FABACEAE ichinaceaparadoxa\/a(.paradoxa(J.B.S.Norton) AmorpliaouachitensisWWbuf[12]both;shrub Britt,[7]Oz;perennialforb FAGACEAE LiatrAis.Gsrqauyan|8o]sOau(;L.)peMriecnhnxi.avlarf,orrbo;mpcceinrtfearToofrrd,is&- QuerOcuu;satcreerei;folailat(hPoaulmgehr)lHoensgs&thSotouygnhotfft[o1,1b3e] torfitbhuteioOnuaicnhtihteaNMovoaucnutlaiitnes.Upliftsubsection cQl.ossheulmyarrdeiliat,erdetcoe,notrreevseeanrcchonisnpdeicciaftiecswtihtaht PolyOmun;iaacnonsusaaltotfoernbs;iskAn.oB.wPnitotnmlaynf&rVo.mBatthees[N1o4-] athrilsiatnasxaonna(isWimloliraemscl2o0s0e3ly|TahffeilgilaotebdalwiptohpuQ-. vMaocuunltiatienUsp.liftsubsectionoftheOuachita lationislessthan600individuals. So//c/ogoouoc/7/tens/sC.&J.Taylor[6]Ou;peren- HYDROPHYLLACEAE nialforb;centerofdistributionintheNova- HydrophyllunibrowneiKrai&Bates[12]Ou;pe- culiteUpliftsubsectionoftheOuachita rennialforb;centerofdistributioninthe Mountains. NovaculiteUpliftsubsectionoftheOuachita l/tYnon/a/effermonn/Engelm.exA.Gray[12]Ou; Mountains. perennialforb LAMIACEAE BRASSICACEAE Monarda fistulosa ssp. fistulosa L. var. Cardamineangustata0£Sc\-\u\z\tSi[.ouachitano stipitatoglandulosa,comb.nov.ined.[6,8] E.B.Smith[4,5]Ou;perennialforb;known both;perennialforb;thistaxonhasappar- onlyfrorntheNovaculiteUpliftsubsection entlynotbeenvalidlypublishedatthevari- oftheOuachitaMountains. etallevel,althoughWaterfall(1970)firstde- !esquerellafillformisRollins[7]Oz;winterannua scribeditatthespecieslevel,whichwas forb;restrictedtolimestoneglades,almost conceptuallyendorsedbySmith(1988). totallywithintheSpringfieldPlainsubsec- ScutellariabushiiBritt.[7]Oz;perennialforb;re- tionoftheOzarks. strictedtodolomiteglades,withthemajor- StreptanthusmoculatusNutt.ssp.obtusifolius ityoftheworld'spopulationinthedrainages (Hook.)Rollins[8]Ou;annualforb oftheCurrent,ElevenPoint,andWhiterivers. StreptanthussquamiformisGoodman[6]Ou; LILIACEAE annualforb;knownonlyfromtheNovaculite NemastylisnuttalliiPickering[7]Oz;perennial UpliftsubsectionoftheOuachitaMountains. forb CAPRIFOLIACEAE POACEAE ViburnumozarkenseAshe[5,12,14]both;shrub ElymusglaucusBuckleyssp.mackenzii(Bush) COMMELINACEAE J.J.N.Campbell[7,8]both;perennialforb TradescantialongipesE.5.Anderson&Woods. PTERIDACEAE [4,6]both;perennialforb;althoughknown PellaeaglabellaMett.exKuhnssp.missouriensis fromboththeOuachitasandOzarks,this (Gastony)Windham[14]Oz;fern;thisisthe speciesISespeciallycharacteristicinacidic di-ploid,sexualvarietyofawide-ranging woodlandsondissecteduplandsintheCur- species.Exceptforthespores,itismorpho- rentRiverdrainage,andrelativelyrareelse- logicallyindistinguishablefromtheapomic- whereintheInteriorHighlands. ticssp.glabella(Wagneretal.1965). Tradescantiaozarl<anaE.5.Anderson&Woods. [4,5,14]both;perennialforb 1786 BRIT.ORG/SIDA21(3) Table1.continued RANUNCULACEAE SaxifragapalmefiBush[12]both;perennialforb DelphiniumnewtonianumDM.Moore[4,5] SaxifmgavirginiensisMichx.var.subintegra both;perennialforb Goodman[1,1.3]both;perennialforb DelphiniumtreleaseiBushexK.C.Davis[7]Oz; SCROPHULARIACEAE perennialforb AgalinisnuttalliiShinners[17[Ou;annualforb RUBIACEAE PenstemoncobaeaNutt.vai.purpureusPennell GaliumarkansanumA.Gtayvdt.arkansanum[6,8] [7]Oz;perennialforb;althoughthetypical both;perennialforb varietyisawiderangingspeciesoftheprai- GaliumarkansanumA.Grayyar.pubiflorumE.B, riebiome.thisvarietyisrestrictedtoglades Smith[6,8]Ou;perennialforb;morphologi- oncarbonatebedrockintheInteriorHigh- cally,thistaxonappearscloselyrelatedtothe lands. parentvarietyandneedsfurthergenetic VALERIANACEAE eUpvlailfutastiuobns.ecKtnioonwnofotnhleyOfuraocmhitthaeMNoouvnatcauilnist.e Valebrioatnhe;llaanlnounagliffloorrab(Torr.&A.Gray)Walp.[1] HousOtuo;nipaeroeunancihalitfaonrab;(cEe.nB.teSrmiotfhd)isTterrrieblulti[o6n,8i]n Valebrioatnhe;llaannnuutatalllfioirb(Torr.&A.Gray)Walp.[1,8] tchhietaNoMvoaucnutlaiitensU.pliftsubsectionoftheOua- VVaalleerriiaanneellllaapozaolrmkeadnaDyDaylal[1[,71]2]boOtuh;;aannnnuuaallffoorrbb; SAXIFRAGACEAE centerofdistributionintheNovaculiteUp- HeucheravillosaMichx.var.cj/tonsona(Rydberg) liftsubsectionoftheOuachitaMountains. E.B.Smith[12,14]both;perennialforb TheOuachitaMountainscomprise25%oi thetotalareaol theInterior Highlands,butsupport81%oltheendemictaxaoftheInteriorHighlands.Four- teentaxa,representing39%oitheregionsendemicplants,arefoundonlyin theOuachitaMountains.Fifteentaxa(42%)arefoundinboththeOuachitaand Ozarkregions,whileseventaxaQ9%))arerestrictedtotheOzarkPlateau.Also signilicantisthat25%ol theendemicInteriorITighlandslloraisassociated withtheNovaculiteL)j:)liltsubsectionoltheOuachitas.Thisgeologicsubstrate withitsglades,woodlands,andstreamcom]olexcsisanecologicalhotspotlor endemismwithintheInteriorHighlands. Morethanhalf(58%)oftheendeinicspeciesintheInteriorHighlandsare associatedwithgladehabitats(acidic,calcareous,novaculite).Asdefinedby NelsonandLadd(1982),gladesareopenhabitatswithstronglithologiccontrol thataredominatedbyacharacteristicherbaceousvegetation,withsparsetree andshrubcover,shallowsoils,andoftenwithabundantexposedrock.These exposedxericorhydroxcnchabitatshaveextremeenvironmentalparameters includinglong,usuallyannualdroughtperiodsinthegrowingseason,limited waterretentionintheshallowsoils,andintensesolarheating.Manygladesare alsosaturatedthroughmuchofthedormantseason,withfrequentfreeze-thaw cyclesandassociatedsoilupheavals.Droughtandliremaintainmostgladesin anearlytreelessstate. Thelilehistoriesoftheendemicplantsassociatedwithgladesshowdiverse ZOLLNERETAL.,ENDEMICPLANTSOFTHEINTERIORHIGHLANDS 1787 TABLt2.Characteristicplantcommunityaffiliationsof36endemicInteriorHighlandvascularplants. NumbersinthepercentcolumnrefertothepercentofInteriorHighlandsendemictaxaoccurring inthishabitattype.AIIhabitattypeslisted. CommunityComplex Number Percent CentralInteriorHighlandsDryAcidicGladeandBarrens 5 14 OuachitaMontaneOakForest Ozark-OuachitaDryOakWoodland Ozark-OuachitaDry-MesicOakForest 4 11 Ozark-OuachitaMesicHardwoodForest 5 14 Ozark-OuachitaShortleafPine-OakWoodland 8 22 CentralInteriorHighlandsCalcareousGladeandBarrens 8 22 OuachitaNovaculiteGladeandWoodland 8 22 ArkansasValleyPrairieandWoodlands CentralInteriorHighlandsandAppalachianSinkholeand [RepressionPond OuachitaForestedSeep 12 Ozark-OuachitaRiparian 9 25 13 CentralInteriorAcidicCliffsandTalus 2 6 14CentralInteriorCalcareousCliffsandTalus 5 14 15,Ozark-OuachitaFen 16,North-CentralMaple-BasswoodForest 17 South-CentralInteriorLargeFloodplain 1 3 SoutheasternGreatPlainsTallgrassPrairie evolutionarytraitsthatallowforsurvivalinthesehabitats,butwhichpresum- ablyconfernocompetitiveadvantagesinsurroundmgwoodedhabitat.Forex- ample,Echinaceaparadoxavar.paradoxaandLiatrissquarrosavar.compacta areshade-intolerant,long-livedperennialswitheitherdeeprootsorwater-hold- ingsubterraneantissuesandgrowinopenhabitats.TheValerianellaspecies arevernalannualsthatquicklycoiupletetheirlifecyclesinthespringwhen conditionsarerelativelycoolandwet;theyareusuallyassociatedwiththose areasof thegladehabitatthatremainseasonallysaturatedduetowater seepage.Speciesdistributionsmayalsobecontrolledbyanaffinitytoparticu- larsubstrateswithspecificlithologicalcharacteristicandmineralavailability. Forexample,Scutellariahushiiisfoundonlyondolomite;thesegladesarechar- acterizedbyhighlevelsofsolublecationssuchasmagnesiumandcalcium. Twenty-eightpercentoftheendemicfloraoccurinriparianhabitats.Three (Amsoniahubrichtii,Vernonialettermannii,Valerianellapalmeri)oftheten endemicspeciesfoundinriparianareasareassociatedwithglade-likehabitat structures,i.e.rocky,openhabitatsalongephemeralorintermittentstreams. Thesehabitatsarexero-hydric,andmaintainedbyflashystreamflowsandsoil conditionsthatmakethemseasonallyxericandnearlytreeless. Wefoundthat22%oftheendemicspeciesarecharacteristicallyassoci- 1788 BRIT.ORG/SIDA21(3) atcdwithdrypine-oakandoakwoodUmds.ThesepUmtcommunitiesarercki- rix'clyopen,withsparsetomoderatetreecover,adiverseandwell-developed herbaceousunderstorywithaprominentgraminiodcomponent,andarehi- tivelyIrequent(ireregime.Theendemicspeciesassociatedwithwoodlandhabi- tatsappeartobeintolerantofbothextremeexposureanddeepshade.Solidago ouachitensis,\ovexample,appearstorequirefilteredsunlightbutisnotfound inclosedcanopylorestsorthemoreexposedgladehabitats.Woodlandplant communitiesaremaintained inanopencondition bylireandintermittent droughthutha\'cpre\'ailingen\'ironmcntalconditionsrelativelylessextreme anddynamicthantheexposedglades. AcrosstheInteriorHighlandsregionglades,openwoodlands,andinter- mittentstreamsformlandscapecomplexesthatarecloselyassociatedonthe landscape.Theseplantcc~immunitycomplexesarcmaintainedinanopenor treelessconditionbydrought,fire,andflood.Morethan80%oftheendemic tloraoltheInteriorHighlandsoccurintheselandscapecomplexes. Thelistofendemicspresentedmthispaperwillchangeasnewinforma- tionbecomesax-ailableonplantcommunityassociationsandspeciesdistribu- tions,andasnewtaxaarediscoveredandothersrelegatedtosynonymy.Hovv- e\'er,itisnowpossibletocomparetheInteriorHighlandsregionwithadjacent regionsthatarephysiographicallydifferent.Forexample,the36endemicvas- culartaxaol theInteriorHighlandsrepresentperhaps2%ofthenativeflora, butexhibitnopatternsofendemismatorabovethegenuslevel.Infact,the InteriorHighlandsforallofitsphysiographicuniqueness,includingage,long- termisolationiromitsmoiety—theAppalachianRegion—anditsreputationas a"well-knownrefugium"(Meyerf997)showssurprisingly littlefloristic uniquityIromsurroundingregions.Bycomparison,theWestGulfCoastalPlain withvirttiallynoreliefandnoantiquity,hasaboutfOOendemictaxa(about threepercentofitsflora)ofwhichthreeareabovethespecieslevel(MacRoberts etal.2002).Itisnotknownw^hetherthissamepatternandlevelofendemism occursamongotherorganismalgroups,buttherearesuggestionsthatendemism ratesmaybehigherthanamongthevascularflora.Forexample,thereare24 endemiccraylishtaxaintheOzarks(TheNatureConservancy2003);thisrepre- sentsmorethansevenpercentofNorthAmericanCrayfishdiversityPrelimi- narydata(Harris&Ladd2003)indicatethattheOzarkregionischaracterized byanunusuallyhighlevelolundescribedlichentaxa,includingatleasttwonew genera,hutcurrentdatagapsprecludedetermininglevelsofendemism. ACKNOWl.HDGMl-NTS Appreciationisextendedtoseveralpeoplewhocontributedinformationorcom- mentstothisproject:TheoWitsell,ArkansasNaturalFleritageCommission, Kaushik Mysorekar,TheNatureConservancy,Phil Hyatt,USForestService, JulianCampbell,TheNatureConservancy,AlanWeakley,UniversityofNorth ZOLLNERETAL,ENDEMICPLANTSOFTHEINTERIORHIGHLANDS 1789 CarolinaBotanicalGarden-Herbarium,ScottSimon,TheNatureConservancy, RobEvans, NatureServe, Ayn Shilsky,The Nature Conservancy,George Yatskievych,MissouriBotanicalGarden,andRonLance,ChimneyRock.We alsothankGaryTucker,StariaVanderpool,andtwoanonymousreviewsfor suggestionsinimprovingthemanuscript. REFERF.NCES Bryant,W.S„W.CMcCoivib,andJ.S.Fralish.1993.Oak-hickoryforests(westernmesophytic/ oak-hickoryforests).In:W.FI.Martin,S.G.Boyce,andA.C.Echternacht,eds.Biodiversity ofthesoutheasternUnitedStates:uplandterrestrialcommunities.JohnWiley,New York.Pp.143-201. Comer,P.,D.Fablr-Langendgen,R.Evans,S.Gawler,C.Josse,G.Kittel,S.Menard,M.Pyne,M.Reid,K. 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