Species conflict at Earth’s edges–Contests, climate, and coveted resources.

PUBLISHED 17October2022

DOI 10.3389/fevo.2022.991714

OPENACCESS

EDITEDBY

R.TerryBowyer, UniversityofAlaskaFairbanks, UnitedStates

REVIEWEDBY

JerichoC.Whiting, BrighamYoungUniversity-Idaho, UnitedStates KevinWhite, Retired,Haines,AK,UnitedStates

*CORRESPONDENCE

JoelBerger joel.berger@wcs.org

SPECIALTYSECTION

Thisarticlewassubmittedto ConservationandRestorationEcology, asectionofthejournal FrontiersinEcologyandEvolution

RECEIVED 11July2022

ACCEPTED 09September2022

PUBLISHED 17October2022

CITATION

BergerJ,BielMandHayesFP(2022)

SpeciesconflictatEarth’sedges–Contests,climate,andcoveted resources.

Front.Ecol.Evol. 10:991714. doi: 10.3389/fevo.2022.991714

COPYRIGHT ©2022Berger,BielandHayes.Thisis anopen-accessarticledistributed underthetermsofthe Creative Commons Attribution License (CCBY) Theuse,distributionorreproductionin otherforumsispermitted,provided theoriginalauthor(s)andthecopyright owner(s)arecreditedandthatthe originalpublicationinthisjournalis cited,inaccordancewithaccepted academicpractice.Nouse,distribution orreproductionispermittedwhich doesnotcomplywiththeseterms.

SpeciesconflictatEarth’sedges –Contests,climate,and covetedresources

JoelBerger1,2*,MarkBiel3 andForestP.Hayes1

1 DepartmentofFish,Wildlife,andConservationBiology,ColoradoStateUniversity,FortCollins,CO, UnitedStates, 2 WildlifeConservationSociety–GlobalProgram,TheBronx,NY,UnitedStates, 3 GlacierNationalPark,WestGlacier,MT,UnitedStates

Directconflictbetweenspeciesisaninfrequentlywitnessedbiological phenomenon.Potentialdriversofsuchcontestscanincludeclimate change,especiallyatEarth’shighelevationandlatitudinalextremeswhere temperatureswarm2–5timesfasterthanelsewhereandhydro-geomorphic processessuchasglacialrecessionandsoilerosionaffectspeciesaccess toabioticresources.Weaddressedacomponentofthisbroaderissueby empiricalassessmentsofmammalianconflictoveraccesstofourabiotic resources–minerals,water,snow,andshade–byannotationofpast studiesandbyempiricaldatacollection.EvidenceforNearcticandPalearctic mammalsindicatesthatwhendesertwatersareinshortsupply,contests intensify,generallyfavoringlargerspeciesregardlessoftheirstatusasnative orexotic.Ourempiricaldataindicatethatcontestsbetweentwolarge andapproximatelysimilarly-sizedmammals–mountaingoats(Oreamnos americanus)andbighornsheep(Oviscanadensis)–alonga2,500kmgradient atthreehigh-altitude(abovetree-line)sitesintheRockyMountainsof NorthAmerica,resultinstrikingasymmetries;goatsdominated > 95%of interactions.Despitefarfewerobservationsofencounterstoaccessshade orsnowpatches,anincreasinglyprominentdialogneedstobeheldabout rarelyexploredbiologicalphenomenawherelessisknownthanwemight otherwisepresume,whetherinducedbyclimateorincreasinganthropological alterationbecauseofunderpinningstounderstandcommunitystructure andconservationplanning.Observationsonthefrequencyandintensity bywhichindividualsescalatebehaviortoaccessabioticresourcesremains anunderappreciatedarenatohelpidentifytheproximateimportanceof scarcityinthenaturalenvironment.NotwithstandingDarwin’sprediction some165yearsagothatpopulationsinextremeenvironments(high-latitude, high-altitude)aremorelikelytobeimpactedbyabioticvariablesthanbiotic, conflictbetweenspeciesmaybereflectiveofclimatedegradationcoupled withthechangingnatureofcovetedresources.

KEYWORDS

competition,climatechange,extremes,speciesinteractions,conflict,abiotic resources,mammals,ungulates

Introduction

Ashumans(Homosapiens)continuetheunabated colonizationofEarth’sterrestrialregions(BradshawandBrook, 2014),potentialforconflictoveraccesstorareresources inevitablyincreases.Inextremeenvironments–suchas highlatitudes,theloftiestofelevationsandareasofscant rainfall–abioticforcesmaydictatesurvivalmoredirectly thanspeciesinteractions.Asearlyas1859thiswaspredicted–“WhenwereachtheArcticregions,orsnowcappedsummits,or absolutedeserts,thestruggleforlifeisalmostexclusivelywith theelements”(Darwin, 1859),andsuchlimitationshavebeen amplyconfirmedamongmammalsinArctic,highmountain, anddesertbiomes(Anthony, 1976; CaughleyandGunn, 1993; Daleetal., 1994; Hansenetal., 2019).Nonetheless,persistence underexceptionallyharshconditionsisaboutmorethanabiotic challengeasindividualsmuststillmeetnutrientrequirements andconfigureinteractionswithotherspecies(Krebsetal., 2003; Gauthieretal., 2004; Festa-Bianchetetal., 2011).Whileneither commonnorfrequentlywitnessed,overtcontestsbetween speciesdoensueandmustbeexaminedthroughthebroader lensofglobalchangewhereinteractionsmaybeunmasked particularlyaslandscapeschange.Althoughcompetitionmay takedifferentformsasnotedlongago(Elton, 1946),conflict inextremeenvironmentsshouldnotbediscountedwhere warmingtemperaturesexacerbatesurvivalchallenges(Berger, 2018; Millsetal., 2018).

Alongtheplanet’smostnorthernandsouthernedgesand atthehighestaltitudestemperatureswarm2–5timesfaster thanelsewhere(Pörtneretal., 2021).Consequently,theworld’s mountainsareexperiencingmassglaciallossesfomentedby anaccentuationinthetimingandintensityofwaterflow (Barnettetal., 2005; Leeetal., 2021; Smith, 2021).Suchphysical alterationscreateadditionalcryosphericandgeomorphic changethroughtheredistributionofinorganicinertmaterials byleaching(Butler, 2012; Yangetal., 2021),conversionswhich inturnhaveconsequenceforsoildevelopment,minerals,and plants(DixonandThorn, 2005; Lambertetal., 2020; Zimmer etal., 2022).Byexample,phosphorusorotherbio-metalscan becomeconcentratedatdepositionalsitesinminerallicks, whichofferessentialmicro-nutrientstogeophagousmammals (Linketal., 2011; Pebsworthetal., 2019).Sodium,inparticular, isaprominentcationinsuchsitesanditplaysaprominentrole inseveralbodyfunctions,includinglactation,thoughasingular universalroleofsodiuminungulatesaltlicksmaynotexist (Kreulen, 1985; Robbins, 1993; Ayotteetal., 2006).

Still,theoverarchingimportanceofaccesstoarestricted abioticresourcewasnotedasearlyas1741.JohnBartman commentedonthearduousjourneyofwhite-taileddeer (Odocoileusvirginianus):“ thesoil,Isupposecontainssome salineparticlesagreeabletothedeerwhocomemanymilesto oneoftheseplaces”(Seton, 1927).Empiricaldocumentation isnowwidespreadfromenvironsmoreextremethanthe

NewEngland’stemperateforestsofBartman’sexplorations. IntheexceptionallyaridNamibDesert,elephants(Loxodonta africana)travelupto70kmtoaccesswater(Shoshaniand Viljoen, 1992).OnthecomparativelydryTibetanPlateauabove 4,500m,femalewildyaks(Bosgrunniensmutus)seekremnant snowpatchestosustainmilkproductionfornursingoffspring duringwinterwheneveryothersourceofwaterisfrozensolid (Bergeretal., 2015).Mountaingoats(Oreamnosamericanus), aspeciesnotknownforbroadlocomotortravelefficiency (CôtéandFesta-Bianchet, 2003),maycoverupto29km ingeophagouspursuits(Rice, 2010),wheremineralizedsites varyfromcavestoroadsides,outcropsanderodingmountain slopes,somebecauseofglacialattrition,andsoilsbelow trees(CowanandBrink, 1949).Atleastadozenmammalian ordersincludingPrimates,Rodentia,Lagomorpha,Carnivora, Chiroptera,PerissodactylaandArtiodactlyagotogreatlength seekingminerals(Kreulen, 1985; Linketal., 2011; Pebsworth etal., 2019;seealsoabove).

Despitetheseabbreviateddescriptorsofoneroustravel,we knowlittleabouthow,when,orwherecontestsresultingin interspecificcompetitionoccurbetweenspecies,particularly overaccesstofourabioticresources–minerals,shade, water,andsnow.Suchdeficienciesarisesimplybecause overtinterspecificinteractionsarerarelywitnessed.Improving knowledgeinthismostlyunchartedarenaisunderminedby logisticsofdataacquisitionindifficult-to-reachenvironments and,assuredlyhasbeenexacerbatedbyaglobaldeclinein field-orientedscientificinquiry(Ríos-Saldañaetal., 2018).Yet, understandinghowspeciescontestforaccesstoabioticproducts remainsafertileavenueforfuturestudy.

Herewereportonoutcomesofspeciesconflictinmammals fromextremelandscapesconcentratingonpriorityofaccess tothoseabioticresourcesmostlikelytobeaffectedbyclimate challengeandanthropogenicmodification.Specifically,we mobilizedisparateevidenceastohowhydrogeologicaland otheralterationsconflatetoshapeattainmentofabioticrewards (seeschematicin Figure1).Weadopttwoapproaches:(1) contextualizationofthecaseforconflictatacoarsescale throughpreviouslyreportedaggressiveencountersacrossa rangeofgeographies,and(2)presentationofempiricaldata onovertconflictathighelevation(abovetree-line)sitesalong a2,500kmgradientacrosstheRockyMountaincordillera (NorthAmerica).Ourstudysiteswereconcentratedinareas ofrelativelyrecentglacialrecessionwheretwolargeand approximatelysimilarly-sizedmammals–mountaingoats andbighornsheep(Oviscanadensis)contestedforaccessto minerals.Moreover,giventhemagnitudeofrapidchangein globalecologicalcommunitiesduetoanthropogenic-induced impacts,manyofwhichfavorinvasivespeciesfromplantsto fish,andbirdstomammals,weincludedinourassessment ofconflictexotics(i.e.,horse[Equuscaballus],yak),especially becauseofanincreasinglyprominentdialogaboutcurrent andfuturebiodiversityconservationusingecologicalsurrogates

Assumptions,rationale,andframework

Identifyinglimitedabioticresources

Notallresourcesareofequivalentvaluebut,byinference asjudgedbyananimal’sbehavior,theymaybeclassifiedasto desirability.Considersomethinginanimate,anabioticresource likeshade.Mostofuswillhavewitnessedadogorcatseek thermalreliefonahotday,perhapssituatingitselfunderarocky overhangoratree.Humans,otherprimates,andindividualsof manyspeciesdothisofcourse.Insuchscenarioswhereshade isnotlimited,competitionforthesespotswillbeminimal,yet thescientificliteratureoncompetition,whenshadeislimitedfor wildmammals,isscant.Notonlyaredisplacementeventsrarely observed–oratleastnotreported–butasouranthropogenic grasptightens,afocusonitsconsequentimpactstospecies andhowtheyinteractisusefultounderstandcomponents ofglobalchange.

Offundamentalinterestiswhenaresourceisscarceand differentspeciesseektoutilizeitatthesametime.While overtinterspecificencountersmaybefrequentlycircumvented bytemporalseparation(Valeixetal., 2007),orbyatendency toavoidconflictthroughselforopponentassessment(Parker andRubenstein, 1981; Chapinetal., 2019),thefewpapers thatdescribeactivedisplacementssupporttheassumptionthat abioticresourcesareattimesinshortsupply.

Desertwatersofferacaseinpoint(Table1).Nearly300 discreteinterspeciesencountersinvolvingAfricanelephants, rhinos(Dicerosbicornis)andothermammalsatdrinking pointsatthesametimeunderwentalevelofforcedorsubtle displacements(BergerandCunningham, 1998).Aggressive assertationsincludedrushes(orcharges),headthrustsor singularly-directedwalk-approachestowardinterspecifics (Figure2),allofwhichresultedinrapiddisplacements (Table1).Weoperationallycharacterizedthesesortsofabiotic resourcesascovetediftheyweresitesforwhichspecies contestedpriorityofaccess.

Identifyingglaciallossandanthropogenic alterationofmineralizedsites

ChangeswithinourstudyspheresalongRockyMountain cordillera(Figure1)includethoseinduceddirectlyby warmingtemperatures(Martin-MikleandFagre, 2019)and bymoreimmediatebyhumandestructionofhabitat.Higher temperaturesatourthreestudyareas(seebelow)arestrongly associatedwiththephenologyofsnowmeltandplantgrowth, andanupsloperangeshiftofshrubsandtreesintohistorical alpinetundrahabitats.InnorthernMontana,specifically, GlacierNationalPark,85%oftheice/glacialfieldshavebeenlost sincethepark’screationin1916(HallandFagre, 2003).

Theextenttowhichhydrologicalchangescausedby warminghasaffectedminerallicksusedbyungulatesin theRockyMountainsislesscertain.Yet,acrosssegments ofthisbroadregionconstruction,modificationofhighways haveresultedinmassivelossofpreviously-availablehabitat andmineralizedsitesusedbyelk(Cervuselaphus),moose (Alcesalces),mountaingoats,and,undoubtedly,otherspecies

FIGURE1

Representationofhypothesizedpathwaysleadingtohighelevationconflict. (A) Temperature-basedmeltingofglaciersfacilitatesavailabilityof minerallickscoupledwiththeirlosstogeophagousungulatesbyhighwaysconstruction;(right)hydro-geomorphicchangestomineral availability,and (B) 1927photographofexcessivesubterranean-to-surfacewaterflow(GlacierNationalParkarchivephoto).

(Lundgrenetal., 2021).Wefindthatwhenabioticresourcesare inshortsupply,notonlydointerspecificcontestsintensify,but largerdominatespeciesregardlessoftheirstatusasnativeor exotic.

TABLE1 Examplesofinterspeciesconflict,displacementinfivemammalianorders(Artiodactlya,Perissodactyla,Carnivora,Rodentia,andPrimates) overaccesstowaterandmineralswithnotationonpotentialforcontestsforadditionalproducts(shade,snow)including(feral)horsesand (domestic)yaksasexoticspecies.

Dominant

Resource Species* Native

Topography References

WaterGemsbokYesChacmababoonKuisebRiverNamibDesert Hamiltonetal., 19771

HorseNoPronghorn,bighornsheep,muledeerGreatBasinand Coloradodeserts

ElephantYesBuffalo,giraffe,impala,kudu,roanand sableantelope,warthog,waterbuck,plain’s zebra,wildebeest

BlackrhinoYesMountainandplain’szebras,gemsbok, springbok,wildebeest,giraffe,warthog, leopard,lion,brownandspottedhyena, cheetah

Mountainsand basins Berger, 1985; Ostermann-Kelmetal., 2008; Goochetal., 2017; Halletal., 2018

HwangeNPSavannawoodlands

Valeixetal., 2007

EtoshaNPand NamibDesert

Aridsavannasand desert BergerandCunningham, 1998

ShadeBlackrhinoYesGemsbokUniabRiverNamibDesertJBunpublisheddata

GemsbokYesSpringbokDorosCraterNamibDesertJBunpublisheddata SnowNone–seetext

MineralsYakNoTakinJigme-DorjeNPBhutanese Himalayas JBunpublisheddata2

MountaingoatYesBighornsheepGlacierNPRockyMts.Thispaper MountaingoatYesHoarymarmotGlacierNPRockyMts.JBunpublisheddata3

Displacementscausedactivelyorpassivelyasnotedintextwithdominantspeciesconsumingwaterormineralsandsupplantedindividualsofotherspeciesdelayingaccesstoresourceor departingarea.Seealso Jokinenetal. (2014)forspatialoverlapbynortherntemperateungulatesbutwithoutclearcasesofaggression. *Latinnamesprovidedin SupplementaryTable1

1 Commentinpaperbutnodata.

2 Threeencountersin7days:approachbysinglemaleyakcausedmaletakingroup(size=2)andsinglemale(N =2)toreroute.

3 Threeencounters;displacementsoverhumanurine.

(CowanandBrink, 1949; Table2).Thedegreethatlossesof theselowelevationsitesbecauseofhumanconstruction promotedaccesstohighelevationmineralsisnotclearbut accesstosuchmineralizedsitesisnowpossibleinsomeareas becauseicesheetsnolongerexist(HallandFagre, 2003)aswe describe(Figures1, 3).

Materialsandmethods

Weusedtwoapproachestoappraisespeciesinteractions foraccesstofourabioticresources–water,snow,shadeand minerals:(1)asynthesisofpeer-reviewedstudiescoupledwith grayliteratureandopportunisticobservations,and(2)field workacrossthreehighelevationsites.

Assessingconflictandaccesstothree abioticresources–Water,shade,and snow

Webasedourassessmentofdominanceinteractions primarilyondisplacementorobviouscasesofavoidancewhen

membersoftwospeciesapproachedadiscreteabioticresource. Althoughnumerousaccountsarepublished,weexcludedthose unlessinvolvementwasforanobviouscovetedabioticresource. Bywayofexample,muledeer(Odocoileushemionus)crossingan alpinemeadow,whichcausedyellowbelliedmarmots(Marmota flaviventris)toflee(Armitage, 2003),wasnotincludedsince therewasnoindicationthatresourcesusedbythemarmots werethenusurpedbythepassingdeer.Casesofconflictover resources,mostlywater,aretabulatedin Table1

Shade,anotherabioticresource,isnotablyimportantasa thermalrefugeforavarietyofterrestrialvertebratesincluding thatprovidedbycaves,rockyoverhangs,andtrees(Barrettetal., 2004; Pruetz, 2007; Cainetal., 2008).Wewereunabletofind formalreportsofcontestsbetweenspeciesoveraccesstoshade butincludeourlimitedobservations(Table1).

Assessingconflictandaccessto minerals

Ourempirically-basedfieldworkconcentratedatthree sites–theMountEvanregionofColorado(39.5882,–105.6437), theMariasPassareaofGlacierNationalPark,Montana

Examplesofconflictandtolerance.

Closelyrelatedtaxaof mountaingoatsandbighornsheep,herein–femaleandyoungSiberianibex(left)andargalionsamerockyoutcropsintheGobiDesertof Mongolia(R.Reading). (C) BlackrhinoandelephantinEtoshaNationalPark,Namibia(A.Forsyth). (D) Sizeablegroupsofmountaingoatsand bighornsheepatrestsympatrically(andinsetoftheminbroaderlandscape)onCawRidge,Alberta,Canada(F.Dulude-deBroin).

Species* Site

Generallocale Agent Comment References

ElkSelwayRiverCentralIdaho,USAWeatherInverserelationshipbetween useandsoildrying Dalkeetal., 1965

BighornsheepNorquay(lowelevation)Banffarea,Alberta,CanadaRoadconstructionPost-lickdestruction,sheep usedsaltsfromhighway Singer, 1975

MountaingoatMt.WardleKootenayPark,British Columbia,Canada RoadconstructionGravelpitformconstruction enhancedgoatuse

Singer, 1975

Sixungulates1 29minerallicksBanff,Jasper,Yoho,and Kootenay,Canada StatusunclearNeedsupdating2 CowanandBrink, 1949

*CommonandLatinnamesprovidedin SupplementaryTable1

1 Caribou,moose,elk,muledeer,mountaingoats,andbighornsheep.

2 Sitevisitstotheselocalesarenecessarytounderstandcurrentconditions.

(48.3166,–113.3548; Figure3),andCawRidgeinwest-central Alberta(Canada, ∼ 54.8000,–119.8000).Observationswere conductedatGlacierNationalPark(Montana)inlateMay–earlyJunein2020–2021,atMt.Evans,ColoradoJune–July (2020)andJune–August(2021),andatCawRidgein2017–2018 (Dulude-deBroinetal., 2020).

MountaingoatsarenativetotheMontanaandAlbertasites andbighornsheeptoallthree(Festa-BianchetandCôté, 2008). InColorado,however,mountaingoatsarenotnative.Theywere introducedtherein1947andhaveincreasedgreatly,ashasalso beenthecase(alsoasintroducedspecies)inWyoming,Utah, andelsewhere(CôtéandFesta-Bianchet, 2003).

InGlacier,ourobservationsconcentratedatnaturalmineral licks.AtMt.Evans,wefocusedona3,600msitewitheffluent andsaltsoughtbybothgoatsandsheep(Clay, 2019).At CawRidgeconflictswereeitherforaccesstorestingsites orforafewplantmorselsbutnotabioticproducts(see Supplementaryvideos1,2).Despitevariationinelevation andlatitude,theuseofthreehighelevationsitesinthe RockyMountainsofferedanopportunitytogaugewhether thedirectionofdominanceanddisplacementbetweenbighorn sheepandmountaingoatswasconsistent.

Amongthevariablesweconsideredtohaveapossible impactontheoutcomeofencountersweregroupsizes.

FIGURE2

(A) MountaingoatsataminerallickinGlacierNationalPark,MT(W.Sarmento). (B)

TABLE2 Examplesofminerallickusageandchangeinlickusageduetoanthropogenicalterationorlocalweatherconditions.

Generally,itiseasytoenumerateassemblysizebutas individualsbecomemoredispersed,evaluationsgrowmore complex.Forinstance,ahalfdozenclusteredmoosecaneasily becountedasagroupbutifeachindividualisspreadacross severalsemi-distantwillowpatchesit’slessobviousifthisisto beconsideredagroup(MolvarandBowyer, 1994).Different versionsofgroupmetrics(Bowyer, 1987)havebeenreportedfor decades;theseincludeinter-individualdistancesofseparation, cohesiveness,orbehaviorssuchascoordinatedfeedingor resting;littleconsensusexists(Elgar, 1989; Treves, 2000).

Weoperationallydefinedagroupasaclusterofindividuals inwhichthebehaviorofoneislikelytoaffectthatofothers. Oursnapshotapproachwasobviouslyajudgmentbecauseat timestherewascertaintyofresponsebutnotatothertimes.

Forexample,thefivepicturedanimals(atotaloffourgoatsand onesheep)in Figure3B mightbeconsideredonegroupoffive, ortwogroups,respectivelyofoneandfour.Atanintraspecific level,whichistypicallythewaygroupsaredefined,perspective andscalematter,butaswenotebelow,groupsizeshadtrivial,if anyeffect,onoutcomesoveraccesstominerals.

Results

Mostreportsaboutspeciesconflictatabioticresources emanatefrominteractionsobservedatdesertwaters.Bodysize

isnotedasamajordeterminantofoutcome.Elephants,for instance,arenotonlythetypicalvictor,buttheyshowlittle toleranceforotherspecies(Table1).Agonism,dominance,and directionalityislikelytovarybysite,history,andnecessity. Amongferalorotherwiseintroducedspecies,nativemammals maybedelayedordeniedaccess(Halletal., 2018; Ferrettiand Mori, 2020;seealso Table1).

Otherspottilydistributedabioticproductslikeshadeand snowhavereceivedmuchlessscrutinyasresourcesforwhich speciescompetedespitetheirknownbiologicalrelevance (Rosvold, 2016).Beyondwater,however,theonlyevidence forcompetitiontoaccessshadestemsfromobservationsin theNamibDesertorKalahariSandswhereblackrhinos displacedungulatesfromshadetreestheysubsequentlyused (Table1).Withrespecttosnowpatches,wewitnessedsympatry betweenmountaingoatsandbighornsheep,buttheyremained distalwithoutantagonism.Likewise,atabout4,900monthe TibetanPlateauwildyaksandchiru(Pantholopshodgsonii) movedtosnowpatcheswithin200moftheotherwithout apparentcontests.

Unlikesnowandshadewhicharestronglyseasonalfeatures oflocalweather,mineralizedsitesarelesstransientalthough theiravailabilitytoanimalforagersvarieswithinsolation, localhydrology,andsoil.Atminerallicks,mountaingoats dominatedbighornsheepinmorethan95%ofobserved displacements(Figure4).

FIGURE3

Mountaingoat-bighornsheepcontestsatalpinesitesinGlacierNationalPark,MT. (A) Remnantsnowfieldandminerallickwith (B) threegoats (lowerrightcentral)andsinglegoatandsinglefemalebighorntopleftatwater-saturatedminerallick, (C) sympatricfeedingpriortogoat displacementofbighorn(enlarged),and (D) malegoatwithactionablehornthreatdisplacingtwomalebighorn.Insetreflectsfoursitesof knowndisplacementsbetweenmountainsgoatsand(1)Dallsheep(Ovisdalli)inAlaska(Klein, 2019),andbighornsheepat(2)CawRidge, Alberta,Canada(3)GlacierNationalPark,MT,and(4)Mt.Evans,CO.

Regardlessofsite,goatsinitiatedeveryinteraction,andmost involvedpassiveapproaches(73%of106)wherebysubordinate sheepwalkedorskippedaway.In12%ofthetotalcasesneither goatsnorsheephadperceptibleresponses.Rapidapproaches orhornthreats(Figure3D)resultedinflight(< 5%);the longestdistancefledwas ∼ 75 100m.Malegoatsaccounted for68%oftheknownencounters(Figure4).Meangroupsize differedstatisticallybetweenspecies(bighornsheep x = 3 40, SD = 1 44;mountaingoats x = 2 57, SD = 1 43; ttest, p < 0.01)butisunlikelybiologicallyrelevantgiventhefrequencyof overwhelmingdominancebygoats.

Discussion

Interspeciesconflictisnotacommonlydocumented norwell-studiedphenomenon,butitobviouslyoccursas competitionforpatchilyscatteredabioticresources.Likeother rarelyobservedphenomena,suchasinfanticideortooluse, furtherdiscoveryamongwildspecies,awaits.

Bothinterspecificandintraspecificcompetitioncreate knownstrongselectionpressuresthatsculptmorphology, behavior,andecologyviaevolutionarypathways(Mayr, 1982; Bowyer, 2022).Incontemporarysettings,thenatureofconflict toaccessresourcesmayormaynotbechanging,anuncertainty thatexistsbecausewelackecologicalbaselines.Consequently,

weknowlittleaboutwhetherdirectanthropogenicalterations onlandscapeshavefacilitatedanimalmovementsintorealms wherethefrequencyofinteractionshaschanged.While recentclimatealterationsarecreatinghydro-geologicalchange includingofsoilsandthelossofglacialicethataffectthe distributionofabioticresources(Barnettetal., 2005; Leeetal., 2021),duetoaliteraldearthofinformationweknowlittle about if or how theimmediacyofclimatechallengeaffects interspeciescontests.

Atanintraspecificlevel,activecompetitionforabiotic resourcesisknownforreptiles,asupto128Aldabragiant tortoises(Aldabrachelysgigantea)areknowntopileundera singleshadetree(SwinglandandLessells, 1979).Alsoassociated withthermalrefugia,arecasesinvolvingdifferentgenera (Egernia and Eulamprus)ofskinkswhichcompeteforaccessto highelevationcrevasses(LangkildeandShine, 2004).

Nonetheless,theextenttowhichmobilemammalsrelocate toalternativesitesandencounterpossiblegreatercompetition ishighlyuncertainbecauseoftheaforementionedissueofno ecologicalbaseline.Justasitisoftendifficulttogaugepatternsof trendindiseasewhenmonitoringhasbeeninsufficient,similar issuesconfronttheimmediacyofknowingwhetherdirect interspecificinteractionshavechangedinfrequencyacrosstime.

Ourdataoncontestsforabovetreelinemineralsisa caseinpoint.Wedonotknowiftheseminerallicksare anewlydiscoveredresource.Perhapstheyhaveincreasedin

FIGURE4

Outcomesofdominanceassertionbetweenmountaingoatsandbighornsheep. (A) Histograminwhichbluecolumn(goats)is%ofencounters wonrelativetoorange(sheep),bystudysitewiththoseinAlberta,Canadaforspaceorfood,andinColoradoformicro-nutrients. (B) Relative proportionsofgoat-initiateddisplacementsofsheepbytype(asdefinedinMethods),andofknownsex; (C) distributionofgoatandsheep groupsizes.

availabilityduerecenthydro-geomorphicclimate-inducedhigh elevationalterations,asknownfortheHimalayas(Leeetal., 2021),orsomethingelse.Importantlyandregardlessofwhether themediatingforcesaredirectlyhumansuchaslocalhabitat destruction,orbroaderandslowerlikewarmingtemperatures, giventhescarcityandpatchinessofabioticsources(e.g.,mineral licks,desertwaters)opportunitiesforinterspeciesconflictarise overaccess.Clearly,humanalterationsofremotedesertwaters heightentheaccessibilitychallenge(BraithwaiteandMuller, 1997; Simpsonetal., 2011; Larsenetal., 2012),ashasroad constructionwherespecieshavechangedtheirbehaviorto accessminerallicks(Kroesenetal., 2020).

Amonglifehistoryvariablesthatserveasanarbiterof dominanceduringinterspeciesconflictisbodysize(Berger andCunningham, 1998).Withanthropogenicchange,alienmediateddisplacementofnativefaunaoccurs(Berger, 1985; Halletal., 2018)despitescientifically-astutepositionsabout nativityorappropriateecologicalsurrogatesforecological restoration(Lundgrenetal., 2021).Nevertheless,conservation effortsprogressinbothprotectedareasandfurtherafieldby broadeningthedistributionofarid-landswatertoenhance biodiversityandtourism(Simpsonetal., 2011; Larsenetal., 2012).

Conflictbetweenspeciesremainsnotonlyofbroad ecologicalinterestbutharkenstotherootsofscientificcuriosity aboutescalatedaggression.Inthecaseofhighelevation mountaingoats,thespeciesoccupiesabasalpositionin theCapriniclade(ShaferandHall, 2010)withstereotypical canalizedbehaviorassociatedwithprimitivetraitsanda propensityforaggression(Geist, 1971; Festa-Bianchetand Côté, 2008).This,inturn,mayexplainantagonismand dominanceoverbighornsheep(Figure4),asituationwith immediateconservationrelevancegivenrecentcontroversyover introducedmountaingoats.

Asacold-adaptedspecies,mountaingoatssymbolize climatealteration(Whiteetal., 2018),aresoughtbyvisitors inplaceslikeGlacierNationalPark,andremainemblematical fortheGreatNorthernPacificRailway.Thespecieswas introducedtosouthernlocalesaspreviouslynoted,andpublic andscientificopinionsareoftendivisiveaboutsanctityin manyareasincludingGrandTetonandYellowstonenational parks.Recently,nearly60goatswereremovedfromtheformer, actionswithamixofsupport(NationalParkService, 2018) whereasinYellowstonenocontrolsareinplace.Knowledge aboutspeciesdominancetoaccessrareresources,suchas presentedin Figure4,shouldhelpagenciesdeliberateaboutbest conservationpathsforward.

Moreglobally,informationaboutinterspecies contestsamongmammalsremainssparse,asdoes understandingwhethercompetitiveinteractionshavechanged spatiotemporally.Anthropogenicalterationsofthephysical environmentcontinueashabitatsareerasedandasinvasive speciesreworkfoodwebs(Bergeretal., 2020).Whileshade

maybelessapttofunctionasacovetedresource,snowpatches athighelevationdisappearwithindeterminateconsequence (Rosvold, 2016).Bycontrast,waterandmineralsareclearly soughtwherespeciesengageforaccess.Thatweunderstand littleoftheprocesscomplexitythatunderminesproduction andchangeinavailabilityofmanyabioticproducts,otherthan theheightedpaceofhuman-wreakedlandscapemodifications, suggestsafertilegroundforfuture insitu fieldinquiriesabout speciesinteractions.

WebeganthisnarrativepointingtoDarwin’sprescience thatabioticprocessesmaybepotentiallymoredemographically limitingthanbioticfactorsinextremeareas.Wenowknow highlatitudeandhighelevationssitesarechangingmore rapidlythanelsewhereduetoclimate,butmorepopulated areasarelosinghabitatsmorerapidly(Caroetal., 2022).How thesefactorsaffectcommunitiesasspeciesaremoreoften broughttogether,resultinginincreasedcompetitionforabiotic resources,isunclear.Ifconservationpractitionersconsiderthat changesinabioticresourceavailabilitycanincreasecompetition, preemptivemanagementdecisionscanbeimproveduponto facilitatedesirableoutcomes.Forexample,justaslimitedwaters canbemanagedtobenefitbiodiversity,proactiverecognition, andmanagementofexistingandemergingminerallicks mayfacilitateconservationofgeophagousspecies.Moreover, observationsofinterspecificcontestsofferausefulmethodto betterunderstandthelimitingnatureofimportantbiological elements.If–whichisanimportantcaveat–underglobal change,contestsforabioticvariablesincrease,thiscould beanadditionalunforeseenconsequenceofclimatechange withimpactstobiodiversity.If,however,curiosityabout naturecoupledwithfieldstudiescontinuetheirdecline,we willneverknow.

Dataavailabilitystatement

Therawdatasupportingtheconclusionsofthisarticlewill bemadeavailablebytheauthors,withoutunduereservation.

Ethicsstatement

TheanimalstudywasreviewedandapprovedbyIACUC, ColoradoStateUniversity.

Authorcontributions

JBconceivedtheideasofthismanuscriptthroughprolonged conversationswithFPHandMB.Allauthorscontributedtothe fieldwork,aswasthewriting.

Funding

FinancialsupportprovidedbytheWildlifeConservation Society,ColoradoStateUniversity,GlacierNationalPark,and GlacierNationalParkConservancy.

Acknowledgments

Weappreciatethewhole-heartedflexibilityofourfunders todevelopinsightsintohownatureworksandconsiderfuture conservationavenues.WeareindebtedtoF.Dulude-deBroin forprovidingobservationaldata,sharingbiologicalinsights, andprovidingfeedbackonthismanuscriptbasedonthelongtermprojectatCawRidge,Alberta,Canada.Wearegrateful toJ.Lambertforcriticalquestionsandstrongediting,and toS.EkernasandDenverZoologicalSocietyforinsightson competitiveinteractionsfromMtEvans,Colorado.Wefurther appreciateDenverMountainParksandGlacierNationalPark Conservancyforfacilitatingfieldresearchandprovidingpartial funding.WearethankfultoJ.Harringtonforprovidingtwo additionalobservationsofinteractionsatMtEvans,andthe gracioususeofphotosbyF.Dulude-deBroin,R.ReadingandA. Forsyth.WealsothankR.T.Bowyer,J.C.Whiting,andK.White forprovidingvaluablesuggestionstoimprovethismanuscript.

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