WildHorsesasNativeNorthAmericanWildlife

byJayF.Kirkpatrick,Ph.D.andPatriciaM.Fazio,Ph.D. (RevisedJanuary2010)
20032010,Drs.JayF.KirkpatrickandPatriciaM.Fazio.AllRightsReserved.

Arewildhorsestrulywild,asanindigenousspeciesinNorthAmerica,or
aretheyferalweedsbarnyardescapees,farremovedgeneticallyfromtheir prehistoricancestors?Thequestionathandis,therefore,whetherornot modernhorses,Equuscaballus,shouldbeconsiderednativewildlife. Thequestionislegitimate,andtheanswerimportant.InNorthAmerica,the wildhorseisoftenlabeledasanonnative,orevenanexoticspecies,bymost federalorstateagenciesdealingwithwildlifemanagement,suchasthe NationalParkService,U.S.FishandWildlifeService,andtheBureauofLand Management.Thelegalmandateformanyoftheseagenciesistoprotect nativewildlifeandpreventnonnativespeciesfromcausingharmfuleffects onthegeneralecologyoftheland.Thus,managementisoftendirectedattotal eradication,oratleastminimalnumbers.Iftheideathatwildhorseswere, indeed,nativewildlife,agreatmanycurrentmanagementapproachesmight becompromised.Thus,therationaleforexaminingthisproposition,thatthe horseisanativeornonnativespecies,issignificant. ThegenusEquus,whichincludesmodernhorses,zebras,andasses,istheonly survivinggenusinaoncediversefamilyofhorsesthatincluded27genera. TheprecisedateoforiginforthegenusEquusisunknown,butevidence documentsthedispersalofEquusfromNorthAmericatoEurasia approximately23millionyearsagoandapossibleoriginatabout3.43.9 millionyearsago.Followingthisoriginalemigration,severalextinctions occurredinNorthAmerica,withadditionalmigrationstoAsia(presumably acrosstheBeringLandBridge),andreturnmigrationsbacktoNorthAmerica, overtime.ThelastNorthAmericanextinctionprobablyoccurredbetween 13,000and11,000yearsago(Fazio1995),althoughmorerecentextinctionsfor horseshavebeensuggested.Dr.RossMacPhee,CuratorofMammalogyatthe AmericanMuseumofNaturalHistory,andcolleagues,havedatedthe existenceofwoollymammothsandhorsesinNorthAmericatoasrecentas 7,600yearsago.Haditnotbeenforpreviouswestwardmigration,overthe

2 BeringLandBridge,intonorthwesternRussia(Siberia)andAsia,thehorse wouldhavefacedcompleteextinction.However,Equussurvivedandspreadto allcontinentsoftheglobe,exceptAustraliaandAntarctica. In1493,onColumbussecondvoyagetotheAmericas,Spanishhorses, representingE.caballus,werebroughtbacktoNorthAmerica,firstinthe VirginIslands,and,in1519,theywerereintroducedonthecontinent,in moderndayMexico,fromwheretheyradiatedthroughouttheAmericanGreat Plains,afterescapefromtheirownersorbypilfering(Fazio1995). CriticsoftheideathattheNorthAmericanwildhorseisanativeanimal,using onlyselectedpaleontologicaldata,assertthatthespecies,E.caballus(orthe caballoidhorse),whichwasintroducedin1519,wasadifferentspeciesfrom thatwhichdisappearedbetween13,00011,000yearsbefore.Hereinliesthe cruxofthedebate.However,neitherpaleontologicalopinionnormodern moleculargeneticssupportthecontentionthatthemodernhorseinNorth Americaisnonnative. Equus,amonophyletictaxon,isfirstrepresentedintheNorthAmericanfossil recordaboutfourmillionyearsagobyE.simplicidens,andthisspeciesis directlyancestraltolaterBlancanspeciesaboutthreemillionyearsago (AzaroliandVoorhies1990).Azzaroli(1992)believed,againonthebasisof fossilrecords,thatE.simplicidensgaverisetothelatePlioceneE.Idahoensis, andthatspecies,inturn,gaverisetothefirstcaballoidhorsestwomillion yearsagoinNorthAmerica.SomemigratedtoAsiaaboutonemillionyears ago,whileothers,suchasE.niobrarensis,remainedinNorthAmerica. InNorthAmerica,thedivergenceofE.caballusintovariousecomorphotypes (breeds)includedE.caballusmexicanus,ortheAmericanPeriglacialHorse (alsoknownasE.caballuslaurentiusHay,ormidlandensisQuinn)(Hibbard 1955).Today,wewouldrecognizetheselattertwohorsesasbreeds,butinthe realmofwildlife,thetermusedissubspecies.Byecomorphotype,wereferto differingphenotypicorphysicalcharacteristicswithinthesamespecies, causedbygeneticisolationindiscretehabitats.InNorthAmerica,isolated lowermolarteethandamandiblefromsitesoftheIrvingtonianageappearto beE.caballus,morphologically.ThroughmostofthePleistoceneEpochin NorthAmerica,thecommonestspeciesofEquuswerenotcaballinesbutother lineages(species)resemblingzebras,hemiones,andpossiblyasses(McGrew 1944;Quinn,1957).

3 InitiallyrareinNorthAmerica,caballoidhorseswereassociatedwithstenoid horses(perhapsancestralforerunnersbutcertainlydistinctspecies),but betweenonemillionand500,000yearsago,thecaballoidhorsesreplacedthe stenoidhorsesbecauseofclimaticpreferencesandchangesinecological niches(Forstn1988).BythelatePleistocene,theNorthAmericantaxathatcan definitelybeassignedtoE.caballusareE.caballusalaskae(Azzaroli1995)and E.caballusmexicanus(Winans1989usingthenamelaurentius).Both subspecieswerethoughttohavebeenderivedfromE.niobrarensis(Azzaroli 1995). Thus,basedonagreatdealofpaleontologicaldata,theoriginofE.caballusis thoughttobeabouttwomillionyearsago,anditoriginatedinNorthAmerica. However,thedeterminationofspeciesdivergencebasedonphenotypeisat leastmodestlysubjectiveandoftenfailstoaccountforthediffering ecomorphotypeswithinaspecies,describedabove.Purelytaxonomic methodologieslookedatphysicalformforclassifyinganimalsandplants, relyingonvisualobservationsofphysicalcharacteristics.Whileearlier taxonomiststriedtodealwiththesubjectivityofchoosingcharacterstheyfelt wouldadequatelydescribe,andthusgroup,generaandspecies,these observationswerelackinginprecision.Nevertheless,themoresubjective paleontologicaldatastronglysuggeststheoriginofE.caballussomewhere betweenoneandtwomillionyearsago. Reclassificationsarenowtakingplace,basedonthepowerandobjectivityof molecularbiology.Ifoneconsidersprimateevolution,forexample,the molecularbiologistshaveprovideduswithacompletelydifferent evolutionarypathwayforhumans,andtheyhavedescribedentirelydifferent relationshipswithotherprimates.Noneofthiswouldhavebeenpossible priortothemethodologiesnowavailablethroughmitochondrialDNA analysis. Aseriesofgeneticanalyses,carriedoutattheSanDiegoZoosCenterfor ReproductioninEndangeredSpecies,andbasedonchromosomedifferences (Benirschkeetal.1965)andmitochondrialgenes(GeorgeandRyder1986)both indicatesignificantgeneticdivergenceamongseveralformsofwildE. caballusasearlyas200,000300,000yearsago.Thesestudiesdonotspeakto theoriginsofE.caballusperse,buttheydopointtoagreatdealofgenetic divergenceamongmembersofE.caballusby200,000to300,000yearsago. Thus,theoriginhadtobeearlier,but,attheveryleast,wellbeforethe disappearanceofthehorseinNorthAmericabetween13,00011,000yearsago.

4 Therelativelynew(30yearold)fieldofmolecularbiology,using mitochondrialDNAanalysis,hasrecentlyrevealedthatthemodernor caballinehorse,E.caballus,isgeneticallyequivalenttoE.lambei,ahorse, accordingtofossilrecords,thatrepresentedthemostrecentEquusspeciesin NorthAmericapriortoextinction.NotonlyisE.caballusgenetically equivalenttoE.lambei,butnoevidenceexistsfortheoriginofE.caballus anywhereexceptNorthAmerica(Forstn1992). AccordingtotheworkofresearchersfromUppsalaUniversityofthe DepartmentofEvolutionaryBiology(Forstn1992),thedateoforigin,based onmutationratesformitochondrialDNA,forE.caballus,issetat approximately1.7millionyearsagoinNorthAmerica.This,ofcourse,isvery close,geologicallyspeaking,tothe12millionyearfigurepresentedbythe interpretationofthefossilrecord. CarlesVil,alsooftheDepartmentofEvolutionaryBiologyatUppsala University,hascorroboratedForstnswork.Viletal.(2001)haveshownthat theoriginofdomestichorselineageswasextremelywidespread,overtimeand geography,andsupportstheexistenceofthecaballoidhorseinNorth Americanbeforeitsdisappearance,corroboratingtheworkofBenirschkeetal. (1965),GeorgeandRyder(1995),andHibbard(1955). AstudyconductedattheAncientBiomoleculesCentreofOxfordUniversity (Weinstocketal.2005)alsocorroboratestheconclusionsofForstn(1992). DespiteagreatdealofvariabilityinthesizeofthePleistoceneequidsfrom differinglocations(mostlyecomorphotypes),theDNAevidencestrongly suggeststhatallofthelargeandsmallcaballinesamplesbelongedtothesame species.Theauthorstates,Thepresenceofamorphologicallyvariable caballinespecieswidelydistributedbothnorthandsouthoftheNorth Americanicesheetsraisesthetantalizingpossibilitythat,inspiteofmanytaxa namedonmorphologicalgrounds,mostorevenallNorthAmericancaballines weremembersofthesamespecies. Inanotherstudy,Krugeretal.(2005),usingmicrosatellitedata,confirmsthe workofForstn(1992)butgivesawiderrangefortheemergenceofthe caballoidhorse,of0.86to2.3millionyearsago.Atthelatest,however,that stillplacesthecaballoidhorseinNorthAmerica860,000yearsago.

5 TheworkofHofreiteretal.(2001),examiningthegeneticsofthesocalledE. lambeifromthepermafrostofAlaska,foundthatthevariationwaswithinthat ofmodernhorses,whichtranslatesintoE.lambeiactuallybeingE.caballus, genetically.Themolecularbiologyevidenceisincontrovertibleand indisputable,butitisalsosupportedbytheinterpretationofthefossilrecord, aswell. Finally,veryrecentwork(Orlandoetal.2009)thatexaminedtheevolutionary historyofavarietyofnoncaballineequidsacrossfourcontinents,found evidencefortaxonomicoversplittingfromspeciestogenericlevels.This overspittingwasbasedprimarilyonlatePleistocenefossilremainswithout thebenefitofmoleculardata.Acoauthorofthisstudy,Dr.AlanCooper,of theAustralianCentreforAncientDNA,stated,Overall,thenewgenetic resultssuggestthatwehaveunderestimatedhowmuchasinglespeciescan varyovertimeandspace,andmistakenlyassumedmorediversityamong extinctspeciesofmegafauna. Thefactthathorsesweredomesticatedbeforetheywerereintroducedmatters littlefromabiologicalviewpoint.Theyarethesamespeciesthatoriginated here,andwhetherornottheyweredomesticatedisquiteirrelevant. Domesticationalteredlittlebiology,andwecanseethatinthephenomenon calledgoingwild,wherewildhorsesreverttoancientbehavioralpatterns. FeistandMcCullough(1976)dubbedthissocialconservationinhispaperon behaviorpatternsandcommunicationinthePryorMountainwildhorses.The reemergenceofprimitivebehaviors,resemblingthoseoftheplainszebra, indicatedtohimtheshallownessofdomesticationinhorses. Theissueofferalizationandtheuseofthewordferalisahumanconstruct thathaslittlebiologicalmeaningexceptintransitorybehavior,usuallyforced ontheanimalinsomemanner.Considerthisparallel.E.Przewalskii (Mongolianwildhorse)disappearedfromMongoliaahundredyearsago.It hassurvivedsincetheninzoos.Thatisnotdomesticationintheclassicsense, butitiscaptivity,withkeepersprovidingfoodandveterinariansproviding healthcare.Thentheywerereleasedduringthe1990sandnowrepopulate theirnativerangeinMongolia.Aretheyareintroducednativespeciesornot? AndwhatisthedifferencebetweenthemandE.caballusinNorthAmerica, exceptforthetimeframeanddegreeofcaptivity? Thekeyelementindescribingananimalasanativespeciesis(1)whereit originated;and(2)whetherornotitcoevolvedwithitshabitat.Clearly,E.

6 caballusdidboth,hereinNorthAmerican.Theremightbeargumentsabout breeds,buttherearenoscientificgroundsforargumentsaboutspecies. Thenonnative,feral,andexoticdesignationsgivenbyagenciesarenotmerely reflectionsoftheirfailuretounderstandmodernsciencebutalsoareflection oftheirdesiretopreserveoldwaysofthinkingtokeepalivetheconflict betweenaspecies(wildhorses),withnoeconomicvalueanymore(bylaw), andtheeconomicvalueofcommerciallivestock. Nativestatusforwildhorseswouldplacetheseanimals,underlaw,withina newcategoryformanagementconsiderations.Asaformofwildlife, embeddedwithwildness,ancientbehavioralpatterns,andthemorphology andbiologyofasensitivepreyspecies,theymayfinallybereleasedfromthe livestockgonelooseappellation. Pleaseciteas:Kirkpatrick,J.F.,andP.M.Fazio.RevisedJanuary2010.Wild HorsesasNativeNorthAmericanWildlife.TheScienceandConservationCenter, ZooMontana,Billings.8pages.

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8 Weinstock,J.E.,A.SherWillerslev,W.Tong,S.Y.W.Ho,D.Rubnestein,J. Storer,J.Burns,L.Martin,C.Bravi,A.Prieto,D.Froese,E.Scott,L.Xulong,A. Cooper.2005.Evolution,systematics,andthephylogeographyofPleistocene horsesintheNewWorld:amolecularperspective.PLoSBiology3:17. WinansM.C.1989.AquantitativestudyofNorthAmericanfossilspeciesof thegenusEquus.pp.262297,in:TheEvolutionofPerissodactyles(D.R. ProtheroandR.M.Schoch,eds.).OxfordUniversityPress,NewYork,NY.

JayF.Kirkpatrick,Director,TheScienceandConservationCenter,ZooMontana, Billings,holdsaPh.D.inreproductivephysiologyfromtheCollegeofVeterinary MedicineatCornellUniversity.

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PatriciaM.Fazio,ResearchFellow,TheScienceandConservationCenter, ZooMontana,Billings,holdsaB.S.inagriculture(animalhusbandry/biology)from CornellUniversity,andM.S.andPh.D.degreesinenvironmentalhistoryfromthe UniversityofWyomingandTexasA&MUniversity,CollegeStation,respectively. HerdissertationwasacreationhistoryofthePryorMountainWildHorseRange, Montana/Wyoming.

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