Mountain Lion

14 Pages·2003·1.85 MB·English

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37
Mountain Lion
BeckyM.Pierce
Pumaconcolor VernonC.Bleich
NOMENCLATURE among mountain lions throughout their range. Although the degree
ofsexualdimorphismdidnotvarygeographically,thesizeofmoun-
CN.Mountainlion,puma,panther,catamount,andcougar
tainlionsgenerallyincreasedwithanincreaseinlatitude(Kurten1973;
SN.Pumaconcolor
Iriarteetal.1990).
Thedorsalpelageofamountainlionistawny,andtheygenerally
TAXONOMY arewhiteontheventralsurface(Fig.37.1).Slightvariationincoloris
common,andtheircoatscanhavereddish,yellowish,orgrayishtinges.
Mountainlionshaveacomplicatedtaxonomichistory.Numeroussub-
Therearenoobviouscontrastingmarkingsonthecoatsofadultlions,
species have been described, largely based on morphometric differ-
otherthanblackmarkingsatthebaseofthevibrissaeonthemuzzle,
ences. For example, Young and Goldman (1946) listed a total of 30
onthedorsalsurfaceoftheears,andonthetipofthetail.InLatin,the
subspecies,andCulveretal.(2000)notedthat32namedsubspecies
specificepithetconcolormeans“singlecolor.”
existed. Of those listed by Young and Goldman (1946), at least 13
In contrast to adults, young mountain lions have darker facial
occurred in Canada or the United States (and adjacent Mexico); the
markingsandareheavilyspottedatbirth(Fig.37.2),butthatpattern
remainderhadgeographicdistributionsrestrictedtoMexico,Central
fadesaskittensmature.Thespottedpelageofkittensbecomeslessob-
America,orSouthAmerica.
viousatabout9monthsofage.Bythetimeyoungareapproximately
Themountainlionwasfirstdescribedfromaspecimencollected
2yearsold,thespottinghaslargelydisappeared(Russell1978).The
inBrazil(Marcgrave1648);Linnaeusreclassifiedthemountainlionas
eyesareclosedatbirth,butopenby2weeksofage(YoungandGoldman
Felisconcolorin1771.Jardine(1834)placedthespeciesinthegenus
1946).LoganandSweanor(1999)reportedthatthelightblueeyecolor
Puma,whereitremains(Wozencraft1993).Thehistoricaltaxonomy
ofkittenschangestotheambercolortypicalofadultswhenkittensare
of the mountain lion was reviewed in detail by Young and Goldman
asyoungas5months.
(1946)andCurrier(1983).
Despitethelargedifferenceinbodyweighttypicalofadultmales
Basedonmolecularandmorphologicalinvestigations,themoun-
andfemales,distinguishingthesexesfromadistancecanbedifficult.
tainlionisthoughttohaveevolvedfromanancestorincommonwiththe
Unlessmalesareinanolderagecategory,withwell-developedshoul-
cheetah(Acinonyxjubatus)andjaguarundi(Herpailurusyaguaroundi)
der,neck,andfacialmusculature,bodyconformationofyoungadult
(Van Valkenburgh et al. 1990; Janczewski et al. 1995; Johnson and
malescanbeconfusedwiththatofolderadultfemales.Genderscan
O’Brien1997;Pecon-SlatteryandO’Brien1998).Mountainlionsare
bedistinguishedbyobservingthegenitalia,butmalelionsfrequently
representedintheNorthAmericanfossilrecorddatingback300,000
are misclassified as females, even by professional wildlife biologists
years(Turner1997).Basedonanextensiveanalysisofmitochondrial
andlawenforcementpersonnel,becausethescrotumandpenisarenot
DNA haplotypes and microsatellite alleles from throughout the geo-
obvioustotheuntrainedobserver.Darkcolorationisassociatedwith
graphicrangeofPumaconcolor,Culveretal.(2000)speculatedthata
hairsurroundingthepenissheathofmales,andisanimportantclueto
massextinctionduringthePleistocene(Martin1989),whicheliminated
thegenderoflions(LoganandSweanor1999).
themajorityoflargemammalsinNorthAmerica,wasfollowedbya
colonizationofmountainlionsofSouthAmericanstock.Furthermore,
Culveretal.(2000)concludedthatmountainlions(northofNicaragua)
todayrepresentasinglesubspecies,Pumaconcolorcouguar,inlieuof
the15thatformerlywererecognized.FiveothersubspeciesofPuma
concolorarerecognizedthroughouttheirsouthernrange(Culveretal.
2000).
DESCRIPTION
Althoughthesexesaresimilarinappearance,maleandfemalemountain
lionsaredimorphicinsize.GayandBest(1995)examinedspecimens
fromthroughouttherangeofmountainlionsandreportedthatmales
weresignificantlylargerthanfemalesin14cranialand5mandibular
measurements.InCalifornia,meanbodyweightofadult(≥2yearsof
age)maleswasX¯=53.4kg(±8.5kg[SD]).Adultfemalesweighed30–
40%less(X¯=35.8±7.7kg)(Charltonetal.1998).Basedonasample
of1076specimensfromOregon,KohlmannandGreen(1999)reported
thatmalesaveragedabout50%heavierthanfemalesofequivalentage.
GayandBest(1995)foundnoevidencethatsexualdimorphismamong
mountain lions varied geographically. Data compiled by Anderson F37.1. Adultmountainlion(Pumaconcolor).S:PhotobyBecky
(1983) show that similar degrees of sexual size dimorphism occur M.Pierce.
744
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MOUNTAINLION(Pumaconcolor) 745
F37.2. Threeweekoldmountainlion(Pumaconcolor).Kittensinthe
birthnursery.S:PhotobyBeckyM.Pierce.
Aswithallfelids,exceptcheetahs,mountainlionshaveretractable
claws(Fig.37.3)attheterminusofeachdigit.Theseclawsfunction
primarily to grasp prey, rather than as an aid in locomotion (Dixon
1982).Therostrumoffelidsisshortandtheorbitsarelarge(Fig.37.4)
comparedtothoseofcanids.Theseskulladaptationsallowforamore
powerfulbite,andfelidsaremoredependentonsightthanarecanids
todetectprey(Vaughanetal.2000).TheadultdentalformulaisI3/3,
C1/1,P3/2,M1/1.
DISTRIBUTION
Althoughtheyoccuratlowdensities,mountainlionsarethemostabun-
dantlargefelidoccupyingNorthAmericaandaresecondinsizeonly
tothejaguar(Felisonca)(Russell1978).Historically,mountainlion
distributionencompassedmostofthewesternhemisphere(Youngand
Goldman1946).TheyoccurredfromtheAtlantictothePacificoceans
andfromapproximately50◦Nto50◦Slatitude,andfromnearsealevel
to about 4000 m elevation (Young and Goldman 1946). In essence,
mountainlionsrangedfromapproximatelytheCassiarRangeofnorth-
ernBritishColumbiatosouthernChileandArgentina,andfromocean
tooceanonthecontinentsofNorthandSouthAmerica.
In North America, the distribution of mountain lions has been
reduced by as much as two thirds of its historical range (Fig. 37.5),
largelybecauseofconflictswithhumansassettlersmigratedwestward
(Rossetal.1997).Currently,mountainlionsoccurinsuitablehabitat
throughoutMexico,inthemajorityofthewesternUnitedStates,and
inwesternCanada;asmallpopulationalsooccursinsouthernFlorida
2 cm
F37.3. Retractileclawmechanismofthemountainlion(Puma F37.4. Skullofthemountainlion(Pumaconcolor).Fromtoptobottom:
concolor).A,Extensorexpansion;B,middleinterphalangealjoint;C,extensor lateralviewofcranium,lateralviewofmandible,dorsalviewofcranium,
tendon;D,flexordigitorumprofundustendon;E,lateraldorsalelastic ventralviewofcranium,dorsalviewofmandible.
ligament;F,distalinterphalangealjoint.S:AdaptedfromGonyeaand
Ashworth(1975).
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746 CARNIVORES
Florida panthers compared to a nearby “population,” and speculated
Kilometers thatdifferencescouldbearesultofgeneralizeddifferencesinhealth
0 400 800 andnutritionalstatusbetweenthetwogroups.Itmayproveusefulfor
Miles mountain lion researchers to examine hematological and serological
0 400 800 data from their study animals, given differences reported within lo-
calizedareasaswellasamongwidelyseparatedpopulations(Dunbar
etal.1997).Forexample,mountainlionsthatusehigh-elevationpasses
tofollowmigratoryprey(Pierceetal.1999b)mayexhibitdifferences
thatarenotevidentinmountainlionsfromthesamepopulation(s)that
donotusehigh-elevationhabitats.Further,dataonhematologicaland
serologicalparametersmayreflectthegeneralhealthstatusofmoun-
tainlionsexistingunderdifferentnutritionalregimes(e.g.,abundantvs.
uncommonprimaryprey)thatmayoccuraslionpopulationsfluctuate
withpopulationsofmuledeer(Odocoileushemionus).
Mountainlions,likemostcats,arewelladaptedtoapproaching
preycloselyandthenrushingforwardquicklytosubduetheirintended
target (Guggisberg 1975). As such, they are not adapted to running
longdistances.Harlowetal.(1992)investigatedtheeffectsofpursuit
onmountainlions.Animalsrespondedphysiologicallytostressesexpe-
riencedduringpursuitbyadepressioninadrenalresponsiveness.They
suggestedthatfrequentpursuitcouldresultindeleteriousphysiologi-
calchangesandurgedcautionbywildlifeagenciesinsettingpursuit
seasonsuntilmoreinformationbecomesavailable.
BEHAVIOR
Activity Patterns. Most felids are nocturnal predators (Kitchener
1991),andmountainlionsaremostactiveduringcrepuscularperiods
Historical distribution (Ackerman1982;Hopkins1989;Sweanor1990).Thatpatternappears
toshifttoamorenocturnalpatternformountainlionsintheproxim-
Present distribution ityofhumandisturbance(VanDykeetal.1986).Mountainlionswill
T37.1. Bloodserumbiochemicalvaluesforfree-ranging
F37.5. Pastandpresentdistributionofthemountainlion(Puma mountainlionsfromCalifornia(n=19)andFlorida(88≤n≤94)
concolor).
Location(X¯±SD)
(Fig.37.5).Recently,mountainlionshavebeenobservedinsoutheast-
ernAlaskaandsouthwesternMinnesota(NowellandJackson1996), Parameter Units California Florida
and they inhabit western North Dakota (Jensen 2001) and Nebraska
(GenowaysandFreeman1996)andarethoughttobecolonizingprevi- Albumin g/dl 3.13a±0.32b 3.70±0.36
Alanineaminotransferase IU/L 58.8±16.7 60.2±35.0
ouslyoccupiedhabitatinOklahoma(Pikeetal.1997).Recentevidence
Alkalinephosphatase IU/L 22.6±11.3 35.4±38.6
ofamountainlioninNewBrunswick,Canada,wasindisputable,butit Aspartateaminotransferase IU/L — 73.4±77.8
couldnotbeascertainedwhethertheanimalhadescapedfromcaptivity Calcium mg/dl 9.53±0.66 9.92±0.66
(CumberlandandDempsey1994).Thepersistenceofmountainlionsin Carbondioxide mEq/L 12.53±1.75 14.33±4.00
themaritimeregionsofCanadaandtheUnitedStatesremainsspecula- Cholesterol mg/dl 155.1±29.9 147.9±26.7
tive(Hansen1992).Anderson(1983)providedadetailedcompilation Chloride mEq/L — 115.5±4.3
of distributional records for mountain lions in the United States and Creatinephosphokinase IU/L — 515.6±415.1
Canada. Creatinine mg/dl 2.05±0.45 1.84±0.54
Gammaglutaminetransferase IU/L — 1.6±1.4
Globulin g/dl 3.45±0.41 —
Glucose mg/dl 110.6±37.3 154.4±51.0
PHYSIOLOGY
Inorganicphosphorus mg/dl 5.66±1.15 5.77±1.51
Therehasbeenapaucityofinformationonbloodparametersforfree- Iron µg/dl — 65.1±33.5
rangingmountainlions.Andersonetal.(1992)soughttorectifythisby Lactatedehydrogenase IU/L — 269.7±173.2
providinghematologicalandbiochemicalreferencevaluesfor≤7adult Potassium mEq/L — 4.60±0.48
Sodium mEq/L — 152.6±3.4
mountainlionsfromColorado.Priortothat,informationwaslimited
Totalbilirubin mg/dl 0.30±0.25 0.26±0.61
toreportsbyCurrierandRussell(1982)orHawkeyandHart(1986).
Totalprotein g/dl 6.58±0.67 7.35±0.67
Paul-Murphyetal.(1994)providedserumbiochemicalreferenceranges Triglycerides mg/dl — 54.9±103.4
for 19 free-ranging mountain lions from throughout California, and Ureanitrogen mg/dl 32.9±6.4 37.7±14.1
concludedthatthevaluesweresimilartothoseindomesticcatsand Uricacid mg/dl — 0.55±0.59
captiveexoticfelids.Dunbaretal.(1997)describedhematologicaland
serumbiochemicalvaluesformountainlionsfromFlorida.Mostvalues S:DatahavebeenadaptedfromPaul-Murphyetal.(1994)andare
wereconsistentwiththoseestablishedformountainlionsinColorado reporteddirectlyfromDunbaretal.(1997).
aThenumberofdigitstotherightofthedecimalpointhasbeenreduced,
(CurrierandRussell1982)andCalifornia(Paul-Murphyetal.1994),
byrounding,forconsistencywiththedatareportedbyDunbaretal.(1997)for
butsomedifferenceswereapparent(Table37.1).
mountainlionsfromFlorida.
Dunbaretal.(1997)attributedhigherpackedcellvolume,hemo- bPaul-Murphyetal.(1994)reportedmeansandtwostandarddeviationsfor
globin,andredbloodcellvaluesreportedbyCurrierandRussell(1982) serumbiochemistryvalues.Standarddeviationsreportedherewerecalculated
tothehigherelevationsinColoradothaninFlorida.Theyalsoreported byhalving,androundingasappropriate,datafromPaul-Murphyetal.(1994)
higher values for these three parameters within one “population” of formountainlionsfromCalifornia.
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MOUNTAINLION(Pumaconcolor) 747
repeatedly move and wait in ambush during the night when hunting houndsforresearchdoesnotappeartocausechangesinhomerangeuse
(Beieretal.1995),butthatbehaviorissuspendedwhencachedprey orabandonmentofpreycachesinmostinstances(Seidenstickeretal.
isavailable(Beieretal.1995;Pierceetal.1998).Movementtoprey 1973;Maehr1997;Pierceetal.1998,1999b).Mountainlionsremain
caches at night occurs earlier for females with small young than for closetohumanactivitydirectlyafterbeingpursued,captured,andthen
othersocialcategoriesofmountainlions(Pierceetal.1998).Thatbe- releasedforresearchpurposes(Seidenstickeretal.1973;Beieretal.
haviorcouldbeastrategytoavoidadultmales,aresponsetogreater 1995).However,theyalsomayavoidareaswheretheyarerepeatedly
energeticneedsofthemother,oratendencytokillpreyclosertothe harassed(HebertandLay1997;JanisandClark2002).
daytimerestinglocationoftheyoung.Femalemountainlionswithout Mountainlionsmovequietlyandlowtothegroundwhenhunt-
youngcovermoreareaandmakelongermovementsthanfemaleswith ingandstalkingprey(Koford1946;Wilson1984;BankandFranklin
young(Sweanor1990).Malesmakethelongestmovements(Sweanor 1998).Earsalsomaybeloweredtoreducedetectionwhiletheymove
1990; Beier et al. 1995), cover the greatest area (Seidensticker et al. slowlyanddeliberately,frequentlyfreezinginpositionwhilewaiting
1973;Loganetal.1986a;Pierceetal.1998),andaremorelikelyto toapproachcloselyenoughtoambushtheintendedprey(Leyhausen
leaveapreycachebeforeitiscompletelyconsumed(Ross1994).The 1979). When prey is within a suitable distance, they rush at the tar-
moreextensivemovementsofmaleslikelyarenecessarytomonitoras get.Pursuitofpreybymountainlionsisrelativelyshortcomparedto
manyfemalesaspossibleandresultinlargehomeranges. thelongchasesofcoursingpredators,suchaswolves(Canislupus).If
Adult mountain lions spend a majority of time resting during a preyiscaught,itusuallyisattackedatorneartheneck.Whenattack-
diel cycle (Beier et al. 1995). The long periods of rest exhibited by ingungulates,mountainlionsmostfrequentlygriptheanteriorventral
mostfelidsmayreduceenergyexpenditureforaspeciesthatoftenkills portionoftheneck,deliveringacrushingbitetothetracheaandcaus-
prey as large as or larger than itself and frequently has prey caches ingsuffocation(Kitchener1991).Thedorsalportionoftheneckalso
available.Mountainlionswillgorgewhenalargeamountoffoodis maybebitten,andthespinalcordcanbesevered.Afterkillingprey,
available(DanvirandLindzey1981;Ross1994)andmovemorefre- it is usual for mountain lions to open the body cavity of the carcass
quentlywhentheyareunsuccessfulatmakingakill(Beieretal.1995). andremovethedigestivetract(Hornocker1970).Otherinternalorgans
Mountainlionsalsoinhabitregionsofextremetemperatures:resting includingtheheart,liver,andlungsoftenareconsumedfirst(Danvir
maybethemostefficientwaytopreventoverheatingindesertenviron- andLindzey1981).Gorgingonthemostnutritioustissuesduringthe
mentsorconservingfatstoresinextremelycoldclimates.Thenegative firstdaymaybeanimportantstrategyformountainlionsthatcanlose
relationshipbetweenmass-specificmetabolicrateandbodysizelikely foodtoscavengers(DanvirandLindzey1981)orneedtomovetolocate
influencesthebehaviorpatternsoflargecarnivores(Robbins1993)and mates(Ross1994).
accountsforlongperiodsofinactivitybymountainlions. Inanexaminationofhunter-killedmountainlions,Robinetteetal.
(1959)reportedthat30%hademptystomachs.Mountainlionsmaycon-
Communication. Mountain lions are the largest felid that make the
sume>10kgofmeatduringasinglefeeding(Hornocker1970;Danvir
low rumbling noise called a “purr.” Purring, chirping, and whistling
andLindzey1981;Ackermanetal.1986),withmorebeingconsumed
vocalizationshavebeendescribedforadultfemaleswhentheyreturn
duringthefirstdaythanonconsecutivedaysfollowingthekill(Danvir
totheiroffspring(Loganetal.1996).Defensivenessandaggressionare
and Lindzey 1981). Results of field sampling and trials with captive
expressedbyasuiteofsimilarbehaviorssharedamongfelids(Kitchener
animalssuggestthatmountainlionsconsumeanaverageof4.4kgof
1991):earslayeddowninaflattenedposition,growling,openingthe
meat/day(DanvirandLindzey1981;Ackermanetal.1986).Inmost
mouth wide while making a hissing sound, tail twitching, and pilo-
instances,preycarcassesaredragged,sometimes>200m,tolocations
erection.Femalemountainlionsinestrusmakealoud“caterwauling”
withvegetativeortopographiccover(Beieretal.1995).Whenlions
vocalizationtoadvertisetheirconditiontomales,andthisbehaviorcon-
finishfeeding,theyusuallyconcealthepreybycoveringitwithsticks,
tinuesthroughoutanyresultingmatingassociation(Rabb1959;Padley
grass,andothermaterialuntilthecarcassisconsumedorabandoned.
1991;Beieretal.1995;Loganetal.1996).
Mountainlionsnormallyconsume73–79%ofadeercarcass(Danvir
Homerangeareasofmountainlionsaremarkedbyscrapesmade
andLindzey1981;Ackerman1982).Coveringcachesmayslowdecom-
bypawingthegroundinabackwardmotion,drawingdirtandground
positionofthemeatandhideitfromscavengers(Beieretal.1995).Prey
litterintoasmallpilebehindtwoparallelgrooves(Seidenstickeretal.
cachesmaybeguardedtoprotectthemfromscavengers,butmountain
1973).Malesmakescrapesmorefrequentlythanfemales(Cunningham
lionscanuseday-bedsseveralkilometersfromkillsites,returningonly
etal.1995),andscrapesoftenhavefecesorurinedepositsassociated
atnighttofeed(Beieretal.1995;Pierceetal.1998,2000b).
with them (Smith 1981; Logan et al. 1996). Scrapes occur most fre-
quentlyinlocationswheretopographyfunnelsanumberofmountain
lionsintothesamearea(Smith1981)andmayfacilitatemutualavoid- REPRODUCTION
ance among individuals with overlapping home ranges (Hornocker
Mountain lions are polygynous and males do not contribute to rear-
1969).MountainlionsdisplayaFlehmanresponsewhensmellingurine
ing young. Like all felids, except African lions (Panthera leo), adult
depositsinscrapes(Seidenstickeretal.1973).Femalesmayscrapeonly
mountainlionsaresolitaryexceptwhenraisingyoung,dispersingwith
wheninestrus(Maehr1997)and,ifso,suchscrapesmaybeusedby
siblings,ormating.Malestendtohaverelativelylargehomeranges,
malestomonitorthereproductivecyclesoffemalesaswellashome
whichoverlapwiththoseofmorethanonefemale(Seidenstickeretal.
rangeoccupancy.Mountainlionsalsoscratchlogsandtreesrepeatedly
1973;Loganetal.1986a;Pierceetal.1999b,2000b).Maleandfemale
withtheirfrontclaws.Scratchmarksmayservethepurposeofmarking
mountainlionslikelyrelyonauditoryandolfactorysignalstolocate
homerangeareas(Schaller1972),butmaysimplybeforsharpening
eachotherformating(Currier1983)(seeBehavior).
claws(Seidenstickeretal.1973).Inaddition,felidshaveglandsinthe
Estrous cycles in females with overlapping home ranges may
cheek,whichoftenarerubbedagainstobjectsandfamilymembersas
besynchronous(Padley1990).Associationsformatinggenerallylast
anothermeansofcommunication(Macdonald1985).
2–5days(Beieretal.1995),duringwhichtimethepairmaycopulateup
Hunting and Feeding. Mountain lions can jump vertically ≥3 m to70timesaday(Eaton1976).Femaleestrouscycleslastapproximately
(Anderson 1983) and are capable tree climbers (Hornocker 1970). 8 days (Rabb 1959; Eaton and Velander 1977) and gestation is 82–
Whenpursuedbycanids,mountainlionsmayseekrefugeintreesor 96 days (Young and Goldman 1946; Anderson 1983; Currier 1983).
rocks(Hornocker1970;Seidenstickeretal.1973;Davisetal.1996). Mountainlionsmayreproduceduringanyseason(Robinetteetal.1961;
Mostreportsofmountainlionsambushingpreyhavedescribedthem Ashmanetal.1983),butmanystudieshaveidentifiedseasonalbirth
stalkingfromtheground(Koford1946;Hornocker1970;Wilson1984; pulses (Robinette et al. 1961; Ashman et al. 1983; Barnhurst and
Beieretal.1995)withveryfewinstancesofattackingpreyfroman Lindzey1984;Loganetal.1986a;Ross1994).Timingofreproduction
elevatedperch(Connolly1949).Theeffectofhuntingbyhumansonthe inmountainlionsmaybeaffectedbyclimateorpreyabundance(Logan
behaviorofmountainlionsisunknown.Pursuitofmountainlionswith etal.1996;Pierceetal.2000a).
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748 CARNIVORES
Femaleshaveeightteats,butonlysixproducemilk(Lechleitner Patagonia(Wilson1984;BankandFranklin1998)andthedesertsof
1969).Thenumberofkittensbornperlittertypicallyrangesfromtwoto thesouthwesternUnitedStates(Cunninghametal.1995;Loganand
four(Robinetteetal.1961;Ashmanetal.1983;RossandJalkotzy1992; Sweanor 2001), by successfully using the limited available cover to
Loganetal.1996),butthismayvarywithpreyavailability,demography, catchprey.Inaddition,Pierce(1999)notedthatmountainlionswere
andpopulationdensity.Femalesmayaveragemoreoffspringperlitter moresuccessfulkillingmuledeerinrelativelyopenhabitatcompared
duringtheirfirstyearofreproductionandfewertowardtheendoftheir tothatinwhichdeerforaged.Thepursuitofdeerbymountainlions
lives(Loganetal.1996).Moststudiesofmountainlionshavereported likelybeganinareaswithvegetativecover,butdeerthatfledintomore
an equal sex ratio among young (Donaldson 1975; Anderson et al. openareaswithoutobstaclesduringthechaseweremorelikelytobe
1992;Loganetal.1996).Padley(1991),however,reportedthatfora caughtthanthosethatremainedinheaviervegetation.
populationofmountainlionsinCalifornia,with12adultfemalesand Mountainlionscanthriveinextremelydryclimateswithlimited
sevenlitters,sixofsevenkittenswhosesexhewasabletodetermine rainfall(Shawetal.1987;Cunninghametal.1995;Loganetal.1996;
were male. In that population, adult males were scarce, and estrous Pierceetal.2000a);however,theseverelycoldtemperaturesofnorthern
females were unable to locate an adult male with which to mate for CanadaandAlaskamaylimittheirdistributiondespitetheavailability
almostayear. ofungulateprey.Mountainlionsmayavoidhumandisturbancewhen
Kittensarebroughttokillsitesbytheirmotherasearlyas8weeks possible(VanDykeetal.1986),butcanpersistnearhumandevelopment
ofage,butarenotweaneduntiltheyare2–3monthsold.Canineteeth (Beieretal.1995;Torresetal.1996).
firstappearbetween20and30daysandmolarsatabout40daysofage. Laing(1988)suggestedthathabitatusebymountainlionsdidnot
Somepermanentteetheruptat5–6monthsofageandpermanentca- differsignificantlyduringa24-hrcycle,butthatpatternmaydepend
ninesappearduringtheeighthmonth,whenprimaryandpermanentca- on the availability of habitats. When habitat used by mountain lions
ninesarepresent.Atweaning,kittensmayweigh3–4kg(Currier1983) fordenningorforcachingpreyisdifferentthanthatusedforresting,
withlittlevariationinweightbetweenthesexesuntilabout30weeks then they will use different habitats during a diel cycle. Beier et al.
ofage(Robinetteetal.1961). (1995)reportedthatmountainlionswithyounghuntedthroughoutthe
Mountainliondensgenerallyarelocatedinrockyterrain(Logan night,butreturnedtothedenduringtheday.Densiteswerelocatedin
etal.1996)orthickvegetation(Beieretal.1995;Bleichetal.1996). nearlyimpenetrablevegetation.Pierceetal.(1998)foundthatmountain
Densarenotelaborate,butdoprovidehidingcover(Ross1994;Beier lionsfedoncachedpreyprimarilyaftersunsetandoftenrestedlong
etal.1995)andthermalbenefits(Bleichetal.1996).Femaleswithkit- distancesfromthecachesiteduringtheday.Inaddition,Beieretal.
tenshaverestrictedmovementpatternsuntiltheyoungareoldenough (1995)reportedthat,insomeinstances,mountainlionsrested>4km
totravelawayfromtheden(Beieretal.1995).Amothermaymove from their cached prey. This behavior also was consistent with the
her young to several different dens before they are weaned (Shaw reported use of high-elevation cliffs during the daytime and use of
1989). lower elevations with flatter terrain for hunting deer at night (Pierce
etal.1998,2000b).
Severalstudieshavedescribedseasonalchangesinelevationand
ECOLOGY
habitatuseassociatedwithchangesinhomerangeformountainlions
Habitat. Thebroaddistributionofmountainlionsatteststotheirability preying on migrating populations of deer and elk (Cervus elaphus)
toadapttoawidevarietyofhabitats.Becauseoftheirwidedistribution (Rasmussen1941;Seidenstickeretal.1973;Ackerman1982;Ashman
(seeFig.37.5),definingpreferredhabitatformountainlionsislimited etal.1983;Murphy1983;Andersonetal.1992;Pierceetal.1999b).In-
togeneralcharacteristicsspecifictodistinctregions,andtremendous deed,differencesinforageavailabilityassociatedwithseasonalranges
variationoccursamongpopulations.Despitethelargerangeofhabitats ofpreyleadstopotentialdifferencesinthetradeoffbetweenforaging
used by mountain lions, relatively few quantitative studies of habitat benefitandpredationriskfordeerandelk.Consequently,theremayex-
selectionexist;manystudiespresentdescriptiveinformationwithout istdifferencesintheeffectsofmountainlionpredationontheseasonal
referencetohabitatavailability.Forthosestudiesthatquantifiedhabitat foragingbehaviorandhabitatselectionofthoselargeungulates(Pierce
selection,preyavailability,vegetation,andtopographyarethegeneral 1999).
characteristicsthatdeterminesuitabilityformountainlions.Withinre-
gionswhereavarietyofhabitattypesoccurs,structureofthevegetation Dispersal. Youngremainwiththeirmotherfor12–18months.Disper-
andtopographyappeartobethemostimportantcriteriafordetermining salmayresultfromthemotherabandoningtheyoung(Seidensticker
habitatuse(Lindzey1987). etal.1973),butitislikelythatthereissomeaggressionbythemother
Most studies of habitat selection for mountain lions in western directedtowardtheyoungtopreventthemfromfollowingher(Hansen
NorthAmericasuggestthatvegetativeandtopographiccover,inaddi- 1992).Dispersalofyoungfromthemotheriscoincidentwiththefemale
tiontosteepslopesandhigherelevations,arepreferredresting,hunting, comingintoestrous;adultmaleswillkillyoung(YoungandGoldman
anddenningsites,whereasopenagriculturallands,sagebrush(Arteme- 1946; Ackerman et al. 1984; Spreadbury 1989; Logan and Sweanor
siatridentata)grasslands,andopenmeadowsandpasturesareavoided 2001).Duringtheperiodofdispersal,mountainlionsareoftenreferred
(Murphy 1983; Logan and Irwin 1985; Laing 1988; Lindzey et al. to as “transients” (Hornocker 1970). Transients, as opposed to “res-
1989).Ashmanetal.(1983)suggestedthatthermalcharacteristicscause idents,” are individuals that do not have a defined home range area
mountain lions to select north-facing slopes at high elevations, with withinapopulation.
morevegetationandcoolertemperatures,inthesummer.South-facing Dispersalisarelativelyriskyperiodduringthelifecycleformany
slopes with little snow cover were selected in winter. Those habitats species (Baker 1978), and survivorship likely declines for mountain
werestronglycorrelatedwiththedensityofdeer.Ruggedorsteepter- lions during the dispersal period (Logan and Sweanor 2001). Tran-
rain,withadequatevegetativecover,maybecriticalforstalkingand sientsaremorelikelythanresidentmountainlionstobeinvolvedin
catchingprey(Ashmanetal.1983;LoganandIrwin1985;Laing1988) depredationincidentsorconflictswithhumans,astheyattempttolo-
orforprovidinghidingcoverandprotectionfromthermalextremesfor catefoodwithouttheadvantageofanestablishedhomerange(Torres
young at denning sites (Belden et al. 1988; Ross 1994; Bleich et al. etal.1996).Malesaremorelikelytodispersefromtheirnatalranges
1996). thanarefemales,andmalestendtodispersefurther(Andersonetal.
Dense hiding cover is preferred by mountain lions for hunting 1992;Sweanoretal.2000).
(Russell 1978; Beier et al. 1995). Indeed, mountain lions were more Mountainlionscandispersenearly500kmfromtheirnatalhome
successfulhuntingpronghorn(Antilocapraamericana)thatinhabited range(LoganandSweanor1999).Consequently,mountainlionhabitat
ruggedterrainwithmorevegetationthanpronghornthatoccurredin throughoutNorthAmericaisconsideredalmostcontiguousonalarge
open prairie habitat (Ockenfels 1994). Mountain lions, however, can scale, with the exception of the isolated populations in southeastern
inhabit open or sparsely vegetated habitats, such as the plateaus of Florida. Nevertheless, mountain lion populations can be isolated by
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MOUNTAINLION(Pumaconcolor) 749
residentialdevelopmentorgeographicfeaturessuchasdesertbasins. FEEDINGHABITS
Reductioninthesizeofhabitat“islands”forpopulationsofanyspecies
Mountainlionskillandeatvertebratepreyalmostexclusively(Lindzey
increasesthechanceofextirpationforthatpopulation(MacArthurand
1987). Vegetation often is ingested because it adhered to a carcass
Wilson1967).
ortohelpwithpassageofparasitesandhairfromthegut(Robinette
Sweanoretal.(2000)describedametapopulationstructure(Levins
et al. 1959; Anderson 1983). The historical distribution of mountain
1970)formountainlionslivinginbasinandrangehabitatsoftheSouth-
lionsthroughoutNorthAmericacoincidedwiththedistributionoftheir
west.Mountainrangesprovideislandsofsuitablehabitatthatsupport
primaryprey—deer.MoststudiesinNorthAmericaidentifieddeeras
semi-isolated populations of mountain lions. Those populations are
the most frequent prey in diets of mountain lions (Hornocker 1970;
maintainedthroughimmigrationbydispersers(primarilymale)from
Shaw1980;Ackerman1982;Loganetal.1996;Rossetal.1997;Pierce
othermountainrangesandrecruits(primarilyfemale)fromwithinthe
etal.2000a).
population.Thebasinsbetweenthemountainsarenotconsideredsuit-
Despitethecongruenceofdeerandmountainliondistributionsin
ablemountainlionhabitat(Germaineetal.2000)andactasbarriersthat
NorthAmerica,lionsaregeneralistpredators(Andersonetal.1992;
maylimitgeneflow.Protectingsourcepopulationsandhabitatstepping
LoganandSweanor1999).Theydopreyonotherlargeungulatesin-
stonesmaybeexceptionallyimportantinconservingsubpopulationsof
cludingmoose(Alcesalces;RossandJalkotzy1996),elk(Hornocker
mountainlionsthatexistinametapopulationstructure(Sweanoretal.
1970),feralhorses(Equuscaballus;Turneretal.1992),feralpigs(Sus
2000). Although concerns about habitat fragmentation and interrup-
scrofa;Harveson1997;Sweitzer1998),andwildsheep(Oviscanaden-
tion of gene flow are legitimate, Ernest et al. (2003) have cautioned
sis; Cronemiller 1948; Harrison 1990; Wehausen 1996; Ross et al.
thatithasnotbeendemonstratedthatmountainlionsconformtothe
1997;Schaeferetal.2000).Mountainlionsalsofeedonsmallerprey
assumptionsinherentinbasicmetapopulationmodels.
suchaslagomorphs(Shaw1980;Ackermanetal.1984);groundsquir-
Survivorship of adults likely varies within most populations on
rels (Spermophilus spp.; Seidensticker et al. 1973); beavers (Castor
anannualbasis(Lindzeyetal.1988),butcanbeaffectedsignificantly
canadensis;Padley1991);porcupines(Erethizondorsatum;Sweitzer
byhunting(Anderson1983).Destructionofhabitatalsocanhavelarge
etal.1997);smallcarnivoresincludingraccoons(Procyonlotor),gray
impactsonthedemographyofmountainlionpopulations.Forexample,
foxes(Urocyoncinereoargenteus),andskunks(Cashmanetal.1992;
fragmentationofhabitatcaninhibitdispersalandincreasecompetition
Beieretal.1995);andbirdssuchaswildturkeys(Meleagrisgallopavo),
forresources(Beieretal.1995;Maehr1997).Furthermore,realestate
grouse,andquail(Ashmanetal.1983;Ackermanetal.1986;Maehr
developmentmaylimitpreypopulations,increasedepredationevents
etal.1990).Currier(1983)listed41nondomesticspeciesofvertebrates
(Shaw1980;Cunninghametal.1995;Torresetal.1996),andincrease
identifiedinthefecesofmountainlionsfromNorthandSouthAmerica.
collisionswithautomobiles(Maehr1997;Beieretal.1995).
Domesticsheep,goats,cattle,horses,dogs,andcatsalsoareconsumed
whenpopulationsofmountainlionsareinproximitytolivestockop-
Disease and Parasites. Anderson (1983) suggested that the widely
erationsorareasofhumanhabitation(Shaw1980;Torresetal.1996).
held opinion that mountain lions are relatively free of parasites and
Attacksonhumansbymountainlionsarerare(Beier1991);however,in
diseases reflected a lack of specific research rather than reality. For
mostinstances,victimsoffatalattacksweretreatedaspreyandeither
example,basedonhisliteraturereview,Anderson(1983)reportedthe
draggedtocoverorpartiallyconsumed(Beier1991;J.Banks,Califor-
occurrence of feline panleukopenia virus was low, but Paul-Murphy
niaDepartmentofFishandGame,pers.commun.,1996).Remainsof
etal.(1994)detectedtiterstothevirusinnearly100%oftheanimals
bobcats(Lynxrufus),coyotes(Canislatrans),andothercompetingcar-
thattheytested.Furthermore,Adaska(1999)reportedthefirstcaseof
nivoresoccasionallyoccurinfecesofmountainlions.Also,speciesthat
coccidioidomycosisdetectedinamountainlion,andJessupetal.(1993)
scavengeonlionkills,includingeaglesandturkeyvultures(Cathartes
documentedthefirstoccurrenceoffelineleukemiavirusinfectionina
aura),canbekilledbymountainlionsguardingapreycache.Although
free-rangingmountainlion.Evermannetal.(1997)reportedontheoc-
intraspecific aggression leading to death in mountain lions may oc-
currenceofpumalentivirusspecificallyinmountainlionsinthestateof
cur as a result of competition for resources or mating opportunities
Washington.Yamamotoetal.(1998)foundoverallseroprevalanceof
(Robinetteetal.1961;Hornocker1970;Sweanor1990;Pierceetal.
bartonellosisin26of74mountainlionstheyexaminedfromCalifor-
1998),instancesofcannibalismbymountainlions,inwhichthecarcass
nia,amongthefirstevidenceofthisconditiondetectedinwildfelids.
hasbeenlargelyconsumed,havebeenreportedfrequently(Lesowski
Hence,itappearsthatAnderson’s(1983)admonitionwasinsightful.
1963;Donaldson1975;Ackermanetal.1984;Spreadbury1989).
Mountain lions potentially are susceptible to many infectious
agentsthataffectdomesticcats(Paul-Murphyetal.1994)aswellasto Selection of Prey. Numerous authors have described the sex, age
otherdiseases(summarizedbyAnderson1983).Someofthese,includ- class, and condition of prey killed by mountain lions. Hornocker
ingfelinepanleukopeniavirus,havethepotentialtoaffectmorbidity (1970),SpaldingandLesowski(1971),Shaw(1977),Ackerman(1982),
and mortality (Paul-Murphy et al. 1994), may limit growth of wild Ackermanetal.(1984),andMurphy(1998)allsuggestedthatvulnera-
populations(Anderson1983),andhaveimplicationsfortheconserva- bilityofindividualpreymaybethemostimportantfactorintheirselec-
tionandpersistenceofaviablepopulationoftheendangeredFlorida tionbymountainlions.Thoseinvestigationsidentifiedindividualsin
panther(Roelkeetal.1993).Foley(1997)concludedthatepizooticdis- youngerorolderageclassesorinpoorerconditionasbeingselectedby
easeslikelyarenotaprimarythreattopopulationsofmountainlionsin mountainlions.Amongadultdeerandbighornsheep,mountainlions
thewesternUnitedStates.Fromahumanhealthperspective,rabieshas alsomaypreyonmalesselectively(Hornocker1970;Ackerman1982;
beendetectedamongmountainlions(Storer1923)andshouldbeof Harrison 1990), especially when males are in a weakened condition
concerninallcasesinvolvingattacksonhumans(Kadeskyetal.1998). followingtherut(Robinetteetal.1959;Shaw1977;Harrison1990)or
Parasitic infections among mountain lions are not uncommon; duringdroughtconditions(LoganandSweanor2001).Studiesofprey
Anderson (1983) listed ≥40 species of parasites that have been col- selection,however,requirecomparisonswithavailabilityofprey.Most
lected.Forresteretal.(1985)andWaid(1990)listedadditionalinter- studiesofpreyselectionbymountainlionshavebeenlimitedbylack
nalparasitesnotnotedbyAnderson(1983).Themajorityofparasites ofinformationonavailabilityofpreyorbybiasesindetectingprey.
recordedhavebeeninternal(protozoans,trematodes,cestodes,ornema- Anderson(1983)attemptedtocomparesexandageratiosofdeer
todes),butmites,ticks,andinsectsareknowntoparasitizemountainli- killedbymountainlionstotheestimatedratiosinthepopulationsof
ons.Penceetal.(1987)notedahighdegreeofoverdispersion(i.e.,afew deerforsixstudyareas,andconcludedthatthesamplingmethodsused
mountainlionsharboredthemajorityofparasitescollectedandmost inthosestudiespreventedcleartestingofanyhypothesisregardingprey
hadfewornoparasites)inhelminthspeciescollectedfromthevisceraof selection.Sincethen,Rossetal.(1997)reportedselectionforyoung
mountainlionsinTexas.Ingeneral,mountainlionshavebeendescribed bighorn sheep by mountain lions. Pierce et al. (2000a) reported that
asremarkablyfreeofexternalparasites(Currier1983),butourexpe- mountain lions selected deer in young and old age classes, and that
riencehasbeenthatticksarecommonlyencounteredduringhandling. femaleswereselectedamongadultdeer.Preferenceforfemalesamong
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750 CARNIVORES
adultmuledeeralsohasbeenreportedforotherpopulationsofmountain pigs(Craig1986)and,inUtah,theykilledtwiceasmanyblack-tailed
lions(BleichandTaylor1998).Morerecentinvestigationssuggested jackrabbits(Lepuscalifornicus)duringwinter(Ackermanetal.1984)
thatmountainlionsdonotselectpreyinpoorcondition(Kunkeletal. thanduringsummer.Adultmaledeermaybetakenatahigherratedur-
1999; Pierce et al. 2000a); however, bone marrow fat has been the ingwinterwhentheyareinweakenedconditionfromtherut(Robinette
primaryindexusedtodeterminepreyconditionbymostinvestigators. etal.1959;Shaw1977).Thevulnerabilityofnewborncalvesresultsin
Althoughpercentagebonemarrowfatwasnotrelatedtopreyselection highdepredationratesoncattleinArizona(Shaw1977;Cunningham
bymountainlions(Kunkeletal.1999;Pierceetal.2000a),theyselected etal.1995).
for older animals. Thus, percentage marrow fat may not adequately
reflect other forms of weakness detected by mountain lions (Pierce
POPULATIONDENSITYANDDYNAMICS
etal.2000a).
Kunkeletal.(1999)determinedthatmountainlionskilleddeer Mountain lions coexist in a system of individual home ranges with
withshorterdiastemaethanthosekilledbywolvesorhunters,butvalues varyingamountsofoverlap(Seidenstickeretal.1973;Hemkeretal.
ofmarrowfatdidnotdifferbetweendeerkilledbymountainlionsand 1984).Meanhomerangesizeanddistributionofmountainlionscanbe
wolves.Sizeofpreyalsoappearstoaffectselectionbymountainlions. affectedbysex(Seidenstickeretal.1973)oravailabilityofresources
Theyselecteddeeroverelk,andelkovermooseinMontana(Kunkel suchasprey(Pierceetal.1999b,2000a).Andersonetal.(1992),how-
etal.1999).RossandJalkotzy(1996)reportedthatmalemountainlions ever, concluded there was no relationship between prey density and
weremorelikelytokillmoosethanwerefemales.Bodysize,however, frequencyofusebymountainlionsofareasinhabitedbydeerandelk.
did not appear to be important for selection of mule deer preyed on Theyfeltthatotherfactors,suchassuitablehabitatforhunting,may
bymountainlionswhencomparedwiththosekilledbycoyotesduring affectpreyavailabilityindependentlyofpreydensity(Kruuk1986)and
winter,whenyoungdeerwereunavailable(Pierceetal.2000a). mayultimatelyaffectthehomerangesizenecessaryformountainlions
Sex and age of mountain lions likely affect their diet. Solitary tobesuccessful.
mountainlionsmaybemorelikelytoeatsmallerpreythandofemales Homerangesofmountainlionsaredelineatedbyvisualandolfac-
withkittens(Ackerman1982).Ackermansuggestedthatkillinglarge torymarkingbehaviors,andadultmalestendtohavelargerhomeranges
prey to provide for their offspring would be a necessary strategy for thanfemales(Seidenstickeretal.1973;Loganetal.1986a;Pierceetal.
mothers,andthatpopulationsofmountainlionscouldnotexistinareas 1998) (Table 37.2) (see Behavior). Male home ranges often overlap
devoidoflargeungulateprey.Pierceetal.(2000a),however,reported thoseofseveralfemalesbuthavelimitedoverlapwiththoseofother
thatfemalemountainlionswithkittens(≤6monthsold)weresignifi- males (Seidensticker et al. 1973; Murphy 1983; Logan and Sweanor
cantlymorelikelytokillyoungdeer(<1yearofage)thanweresingle 2001),whereasthehomerangesofadultfemalesoftenoverlapthose
adultfemalesormales.Birthpulsesofmountainlionpopulationsoften ofotherfemalesextensively(Seidenstickeretal.1973;Sweanoretal.
coincidewiththebirthpulseoftheirprimaryprey(LoganandSweanor 1996;Pierceetal.1999b,2000b;LoganandSweanor2001).Exten-
2001; Pierce et al. 2000a), suggesting that timing of reproduction in siveoverlapamongmalehomeranges(Andersonetal.1992;Sweanor
mountain lions may be dependent on the availability of vulnerable, 1990)andlimitedoverlapoffemalehomeranges(Harrison1990)also
youngprey.Adultfemalesthatarelactatingmaynotbeabletofastfor havebeenreported.Additionally,mountainlionsmayavoideachother
extended periods between unsuccessful attempts to catch larger prey temporarily,abehaviortermed“mutualavoidance”(Hornocker1969),
(Pierceetal.2000a). butnoinvestigationshavetestedforthedominancereversalnecessary
Latitudecorrelatesstronglywithsizeofpreyselectedbymoun- todemonstratetrueterritorialityinmountainlions(Kitchen1974).
tainlions(Iriarteetal.1990;Maehretal.1990).Intemperatezones, Hornocker(1969,1970)andSeidenstickeretal.(1973)suggested
theytendtokilllargerungulates,whereasmountainlionsintropical thatmountainlionpopulationswereselfregulating(seePredator–Prey
environmentshaveahigherfrequencyofsmallpreyintheirdiet.The Dynamics).Theydescribedsocialbehaviorssuchasmutualavoidance,
complicating effect, however, of competition with the jaguar on this territorialmarking,andcannibalismasmechanismsthatlimiteddensi-
patternoflatitudinalpreyvariationisunknown.Additionally,variation tiesofmountainlions.Transientmountainlionswereunabletosecure
inpreyselectionisnotableforsomepopulationsofmountainlionsas permanenthomerangeareasunlessthehomerangeareaofaresident
vulnerabilityoravailabilityofpreychanges.Duringthewetseasonin adultbecamevacant.Thispatternoccurredindependentlyofpreydensi-
coastal California, mountain lions increased predation rates on feral ties.Lindzeyetal.(1994)alsoconcludedthatthedensityofamountain
T37.2. EstimatedhomerangesizeandaveragedensityofmountainlionsfromNorthAmericanstudiesusingtheminimumconvexpolygon
method(Mohr1947)
Males Females
Study Location n Range(km2) n Range(km2) Lions/100km2
RossandJalkotzy1992 Alberta 6 334 21 140 1.9(RA,H),4.2(T)
Cunninghametal.1995 Arizona 5 196 2 109 —(H)
BeierandBarrett1993 California 2 767 12 218 1.1(RA)
Hopkins1989 California 4 199 7 84 3.6(T)a
Andersonetal.1992 Colorado 6 256 7 309 1.1(RA,H)
Maehretal.1991 Florida 8 558 10 191 —
Seidenstickeretal.1973 Idaho 1 453 3 233 0.6(RAb,H)
Ashmanetal.1983 Nevada 7 574 6 178 —(H)
Loganetal.1996 NewMexico 23 187 29 74 1.7(RA)
Murphy1983 Montana 2 462 5 202 7.1(RA,H)
Pittmanetal.2000 Texas 6 349 5 206 0.4(RA,H)
Penceetal.1987 Texas 1 628 5 143 —
Hemkeretal.1984 Utah 1 826 4 685 0.4(T)
Logan1983 Wyoming 2 320 2 73 1.5(RA,H),3.9(T)
N:RA,Densityofresidentadults;T,densityoftotalpopulation;H,huntedpopulation.
aFromAndersonetal.(1992),Table23.
bEstimatedfromSeidenstickeretal.(1973),Table6,Fig.16.
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MOUNTAINLION(Pumaconcolor) 751
lion population in Utah was independent of the density of their pri- independentlyfocusedoneitherthepredatorortheprey,therebylimit-
mary prey, mule deer, because the population of mountain lions did ing conclusions that can be drawn about interspecific effects (Pierce
notincreaseattheratepredictedbytheincreaseinprey.Nonetheless, 1999). In most instances, viable populations of mountain lions are
the population of mountain lions did increase by one third after an dependent on populations of large ungulate prey (Ackerman 1982;
increase in the deer population. Long time lags betweenfluctuations Anderson1983).Mostungulatesthatarepreyedonbymountainlions,
inpreypopulationsandresponsesinmountainlionpopulationsmake includingdeer,elk,andmoose,areaffectedbydensity-dependentpro-
determiningthetruerelationshipbetweenpredatorandpreydifficult cesses(McCullough1979;Boyce1989;Bowyeretal.1999;Keechetal.
(Schaller1972),andemphasizetheneedforlong-termresearchwhen 2000).Therefore,determiningtheeffectofpredationbymountainlions
workingwithlargecarnivores(Pierce1999). onthepopulationdynamicsoftheirpreyrequiresanunderstandingof
To invoke social regulation for a population (Hornocker 1969, thosedensity-dependentprocessesandallfactorspotentiallylimiting
1970;Seidenstickeretal.1973),itmustbedemonstratedthatthepop- thepopulationincludingweather,resources,andotherpredators.
ulationisnotlimitedbycompetitionforresources(WatsonandMoss In the ecological systems that mountain lions inhabit, both ex-
1970).Mutualavoidancedoesnotpreventpassivecompetitionforre- ploitative and interference competition with other predators likely
sources.Pierceetal.(2000b)demonstratedthatcompetitionforpreyby occur. Niches of wolves (Kunkel et al. 1999), coyotes (Pierce et al.
mountainlionsoccurredinapatternthatwouldbeexpectedfornonter- 2000a; Harrison 1990), bears (Murphy 1998), bobcats (Koehler and
ritorialspeciesthatdonotexcludeeachotherfrompreyresources.Fur- Hornocker1991),andjaguars(Iriarteetal.1990)overlapextensively
thermore,ifmountainlionsengagedinaterritorialsystemthatexcluded withthatofmountainlions.Nichepartitioningviahabitatuseandprey
individualsfromresourcesbeyondwhatwouldbeexpectedforapopu- selectionmayallowforcoexistenceofmountainlionswiththoseother
lationlimitedbycompetition,suchcostlybehaviorcouldhaveevolved carnivores.Insomeinstances,scavengingoftheirfoodcachesrequires
onlyifitmaximizedreproductionforindividualsthatwereterritorial mountainlionstoincreasepredationrates(Harrison1990).Inaddition,
and,ultimately,wouldnotleadtopopulationlimitation(McCullough mountainlionsoftenkillsmallerspeciesofcarnivores,suchasbobcats,
1979).Hemkeretal.(1984)concludedthatpopulationsizeofmoun- withoutfeedingonthem(KoehlerandHornocker1991).Conversely,
tainlionswasprimarilylimitedbythepopulationofprey,aconclusion mountainlionshavebeenkilledbypacksofwolves(WhiteandBoyd
supportedbyPierceetal.(2000b)andLoganandSweanor(2001). 1989;BoydandNeale1992).Thoseobservationsalsosuggestthatcom-
Thedensityofmalesmaybelimitedbyintraspecificaggression petitionisanimportantfactoraffectingthepredator–preydynamicsof
to maximize access to females, but mountain lion populations on a multipredatorsystemsthatincludemountainlions.
wholelikelyarenotlimitedbysocialbehavior.Inaddition,dispersalby Mountainlionsareanimportantlimitingfactorforsomeungulate
youngmountainlionsappearstobeindependentofpopulationdensity species(Hornocker1970;Shaw1980).Shaw(1980)estimatedthatlions
(Seidenstickeretal.1973;Hemkeretal.1984;RossandJalkotzy1992; annuallyremoved15–20%ofthemuledeerpopulationontheKaibab
Loganetal.1996),andmountainlionpopulationnumbersparalleltheir PlateauinArizona.Andersonetal.(1992)estimatedthatmountainlions
preypopulations(Lindzeyetal.1994;Loganetal.1996;CoxandStiver annuallykilled8–12%ofthemuledeerpopulationontheUncompah-
1997)(Fig.37.6).Furthermore,LoganandSweanor(2001)reporteda grePlateau,Colorado.InAlberta,asinglemountainlionkilled8.7%
pattern of density-dependent growth for a mountain lion population, (n = 11) of the early-winter bighorn sheep population and 26.1%
providingadditionalevidencethatcompetitionisanimportantfactor (n=6)oftheyoung(Rossetal.1997).
affectingtheirpopulationdynamics. Mountainlionpredationmayreducetheseverityoffluctuationsin
preypopulations(Hornocker1970)byslowingthegrowthrateandre-
ducingovershootsofresource-basedcarryingcapacity(K).Limitation
PREDATOR–PREYDYNAMICS
andregulation,however,aredistinctlydifferentmechanisms.Limitation
Few long-term studies of predator–prey dynamics have been condu- impliesthatthenumberofpreyisreducedthroughdecreasedproduc-
cted that include mountain lions. Unfortunately, many studies have tionorincreasedloss,andthereforeanysourceofpredationislimiting
(Sinclair1989;Boutin1992).Limitationslowstheincreaseofapop-
25,000 25 ulationfromthegrowthrateitwouldhaveachievedintheabsenceof
thepredator.However,ifthepreypopulationcontinuestoincrease,the
proportionofpreyremovedbythepredatorisreduced.Consequently,
20,000 20
preypopulationslimitedbypredatorscancontinuetogrow,although
M
est ou ataslowerratethanintheabsenceofpredation,untiltheyreachK.
v n Regulationimpliesthatrateofremovalbythepredatorchanges
eer har 15,000 15 tain lio watiwthhtihcehpporepdualatotirosnkoilflpprreeyy.;Tashipsrpeyhennuommbeenrosninrcerseualtsse,frsoomdoaensinthcereraastee
e d 10,000 10 n h inthenumberofpredators(numericalresponse),theeasewithwhich
Mul arve (thHeoyllfiinngd1a9n5d9k).ilRlpegreuyla(tfiuonncbtiyonaaplrreedsaptoonrseev)e,notruaalclyomstboipnsatthioeninocfrbeoasthe
s
5,000 5 t ofthepreypopulationandcausesadeclinetolowerdensities,which
ultimatelyresultsinlowerpredationrates.Suchfeedbackmechanisms
cancauseapreypopulationtofluctuatebetweenlow-andhigh-density
0 0 equilibria, both of which are below K. This multiple-equilibria sce-
1930 1940 1950 1960 1970 1980 1990 2000 narioisfrequentlytermeda“predatorpit”(Haber1977;Bergerudetal.
Year 1983;MessierandCrete1985).Functionalresponsesofthepredator
may ultimately determine whether a species has the capacity to reg-
Mule deer harvest ulateapreypopulation(Messier1994).Nonetheless,determiningthe
functionalresponsecurveforaspeciesrequiresanunbiasedmeasureof
Mountain lion harvest predationratethroughouttherangeofpopulationdensitiesoftheprey,
andcontrolledremovalstudiesarenecessarytodeterminethecondi-
F37.6. Harvestofmuledeer(Odocoileushemionus)andmountainlions tions that may cause multiple equilibria (i.e., regulation) to occur in
(Pumaconcolor)inNevada.Muledeerharvestshownincludesmalesonly. predator–preysystems(VanBallenbergheandBallard1994).
Mountainlionharvestshownincludesdepredationremovalsandsporthunting Attemptstomeasurepredationratesofmountainlionshavebeen
(sporthuntingestimatedto1967).S:DatafromS.Stiver,Nevada madeforstaticpopulationsofprey.Usinganenergeticsmodel,Acker-
DepartmentofWildlife,pers.commun.,2001. manetal.(1986)estimatedthatadultmalemountainlionswouldkill
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752 CARNIVORES
adeerevery8–11days,adultfemaleswouldkillevery14–17days,and variable,callingintoquestiontheutilityofthatmeasurement.Despite
afemalewiththreejuvenileswouldkillevery3.3days.Connolly(1949) theproblematicnatureofdiscriminatingamongindividuallions,Beier
reportedthatanadultmountainlionkilledonedeerevery9.7days.Pre- andCunningham(1996)conductedseveraldetailedanalyses.Theyfelt
dationratesonbighornsheepandmuledeerbyfemaleswithkittens itispossibletodetectchangesinrelativeabundanceofmountainlions
ranged from 0.7 to 2.1 ungulates/week (Harrison 1990). Beier et al. usingtrackingtechniquesatthelocallevel.Nonetheless,themigratory
(1995) determined that mountain lions killed 48 large and 58 small natureofsomemountainlionsmakesitproblematictodefinepopula-
mammals/year. Rate of predation by mountain lions can vary exten- tions(Pierceetal.1999b),withseriousimplicationsfortheuseoftrack
sively(RossandJalkotzy1996),andisdependentonanumberoffactors surveysforestimatingchangesinpopulationsize.Tracksurveyscould
includingthenumberandageofyoung(Ackermanetal.1986),season beusefulforestimatingrelativeuseofaspecificgeographicareaby
(Pierce1999),andextentofcompetitionfromotherpredators(Kunkel mountainlions,butinferencesaboutpopulationsizeovertimearecon-
etal.1999)orscavengers(Harrison1990).Thosestudiesgiveinsight foundedbecauseitisdifficulttodefineanopenpopulation.Theuseof
intothebehaviorofmountainlionsandtheirpossibleeffectsonprey tracksurveysoverlargegeographicscalestoindexrelativeabundance
populations,butthequestionofwhethermountainlionscanindepen- ofmountainlions(Smallwood1994;SmallwoodandFitzhugh1995)
dentlyregulateapreypopulationremainsunanswered. hasbeenattempted,butlikelydoesnotprovidethepowernecessaryto
Mountainlionshavebeenimplicatedasregulatingawildhorse detectsuchchanges.
population(Turneretal.1992),andarethoughttohavecausedpopula- Mountain lions frequently are captured to allow attachment of
tionsofporcupinestodeclinethroughpredation(Sweitzeretal.1997). radiotransmitters.Capturetechniquesofteninvolvethepursuitofani-
In addition, some bighorn sheep populations may have declined due malswithhounds,followedbychemicalimmobilization(Loganetal.
to avoidance of high-quality forage in response to predation risk by 1986b;Jessupetal.1986).Physicalretrievalofimmobilizedanimals
mountainlions(Wehausen1996).Inthoseinstances,muledeerwere (Hornocker 1970) is necessary if they are sedated in a location high
theprimarypreyformountainlions.Systemswithmultiplepredators abovetheground.McCownetal.(1990)describedaportablecushion
and multiple prey are more likely to be regulated by predation and thatlessenedtheprobabilityofinjurytomountainlionsfallingfrom
toproducepredatorpits(Gasawayetal.1992).Muledeerpopulations trees.Davisetal.(1996)developedamethodtosafelylowerimmobi-
cankeepmountainlionnumbershigh.Ifpopulationsofalternativeprey lizedmountainlionsfromtreesandcliffs.Footsnareshavebeenused
increase,mountainlionsencounterthosespeciesmorefrequentlyand safelyandsuccessfullybysomeinvestigators(Loganetal.1999;Pierce
predationratesmayincrease,causingalternativepreytodecline.Bleich etal.2000a),andareadvantageousinsomesituations.
etal.(1997)noted,however,thatunlessfemalemountainsheepwere Radiotelemetry has been used extensively to determine the dis-
killedinsignificantproportions,predationbymountainlionsprobably tributionofmountainlionsinnumerousstudies,fromwhichdensity
isnotanimportantfactorinthepopulationdynamicsofthoseungulates. estimateshavebeeninferred.Captureandrecapturemethodsalsohave
Several studies examining the population dynamics of deer and beenusedtoestimatepopulationdensities.Reliabilityofestimatesob-
mountainlionssuggestthatmountainlionsdonotdeterminetheulti- tainedwiththosetechniquesisvariable(LoganandSweanor1999),but
matesizeofdeerpopulations.Instead,otherfactors(especiallyforage densitieshavebeenestimatedat0.3–2.2adultmountainlions/100km2.
resources)appeartobemoreimportantdeterminantsofdeerpopula- Pierce et al. (2000b) estimated the abundance of mountain lions by
tions.Forexample,LoganandSweanor(2001)determinedthatmoun- maintaininganintensiveandconstantefforttocaptureandradio-collar
tain lions were not responsible for the decline of mule deer in New alllionsdetectedonamuledeerwinterrange.Theythenusedaerial
Mexico, but that a lack of forage resources because of drought was telemetrytoestimatethenumberofmountainlionspresentperaerial-
responsible. Pierce et al. (1999a) came to a similar conclusion for a telemetryday,andwereabletodeterminetheirabsoluteabundancefrom
populationofmuledeerintheeasternSierraNevadaofCalifornia.In yeartoyear.Smallwood(1997)notedthatmostvariationindensities
thatstudy,mountainlionnumbersdeclinedprecipitouslyasdidthedeer ofmountainlionscanbeattributedtothespatialextentofindividual
numbers,butwithalagof8years.Longtimelagsbetweenchangesin studyareas,andsuggestedthatfieldstudieswouldbemoremeaningful
preypopulationsandresponsesinmountainlionpopulationsincrease if they spanned larger areas, a variety of land uses and habitats, and
thedifficultyofdeterminingtherelationshipsbetweenmountainlion greaterperiodsoftimethanmostconventionalstudies.
populationsandpreypopulations(Schaller1972),andemphasizethe Moleculartechniqueshavebeendevelopedthatallowinvestigators
needforlong-termstudies(Pierce1999). toidentifyindividualmountainlionsusingDNAobtainedfromblood,
tissue,orhairsamples(Ernest2000a,2000b)andfromfeces(Ernest
etal.2000).Thesetechniquesdependonobtaininggood-qualityDNA,
MANAGEMENTANDCONSERVATION
whichismoredifficultwhenusingfeces(Ernest2000a).FecalDNA
Management Techniques. Demographic information is difficult to hasbeenusedtoidentifyindividuallionsoccurringwithincertainge-
obtainforpopulationsoflarge,crypticpredatorslikemountainlions. ographic areas (Ernest et al. 2000) or associated with particular kill
Tracksurveyshavebeenatraditionalmethodofdeterminingpresence sites for large mammals (Hayes et al. 2000). Use of this technique
orabsenceofmountainlionsaswellasestablishingestimatesofrelative to determine the absolute number of mountain lions occurring in a
abundance.Variousinvestigatorshaveattemptedtodistinguishamong particulargeographicareaissubjecttobiasesassociatedwiththeirdis-
lionsusingtrackcharacteristicsuniquetoindividualanimals(Currier tributionandfrequencyinthearea.Althoughitispossibletoestimate
etal.1977;FitzhughandGorenzel1985;VanDykeetal.1986)ortrack minimumnumbersofindividualspresent,theprobabilityofdetecting
measurements(Currieretal.1977;Shaw1983;FitzhughandGorenzel all individuals using molecular techniques is a function of sampling
1985)torecognizeindividuals.Noneoftheseinvestigatorsbelievedit effort.
wasreliabletodistinguishamongindividualmountainlionsusingonly The literature is replete with estimates for home range sizes of
tracks. mountainlionsthathavebeendeterminedusingland-basedoraerial
Smallwood and Fitzhugh (1993) used a sophisticated statistical telemetry.However,globalpositioningsystem(GPS)technologyraises
technique(multiplegroupdiscriminantanalysis)inanattempttode- questionsregardingpreviousstudies.Bleichetal.(2000)describedthe
velop a method to discriminate among individual animals based on resultsfromGPScollarsontwoadultmalemountainlions.Patternsof
measurementsandcharacteristicsoftracks.However,Grigioneetal. movementwerevastlydifferentandhomerangesweremuchlargerthan
(1999)determinedthatSmallwoodandFitzhugh(1993)hadusedan wereestimatesfromsimultaneousaerialtelemetryflightsconductedon
inappropriatemethod.Basedonadditionalanalyses,theyconcludedit aweeklybasis.SomeGPScollarsincorporateswitches(Bleichetal.
wasunlikelythatindividualmountainlionscouldbedetectedusingdis- 2000)thathavethepotentialtobeusefulinpredictingmountainlion
criminantanalysis.Furthermore,Grigioneetal.(1999)concludedthat activity;Janisetal.(1999)describedandquantifiedtheuseoftraditional
themostcommonlyusedmeasurement(widthofheelpad)fordistin- veryhighfrequencytransmittersequippedwithtipswitchestoestimate
guishingamongmountainlionsbasedontheirtrackswasalsotheleast activityofmountainlions.
P1:JDT
PB336B-37 Feldhammer/0180G July21,2003 17:14
MOUNTAINLION(Pumaconcolor) 753
Translocation of mountain lions as a technique for managing viablepopulationsofmountainlionswillbecomeincreasinglyimpor-
“problem” lions (i.e., animals that have preyed on livestock or that tant(Torres1997).
occupyareasadjacenttohumanhabitation)iscontroversialandisin- Sporthuntingofmountainlionsoccursinthemajorityofstates
consistentwithmanagementpoliciesinsomestatesbecauseofpotential thathaveviablepopulationsofmountainlions,andharvestisamethod
liabilities.Generally,survivaloftranslocatedlionsisverylow.Forex- bywhichwildlifemanagersattempttocontrollionnumbers.Increas-
ample,Ruthetal.(1998)describedanexperimentinwhich14animals ingly,however,theconceptofsporthuntinghascomeundercriticism,
weremovedandreleaseddifferentdistances(X¯ =477km)fromtheir andhuntingofmountainlionshasbeeneliminatedinCalifornia,and
locationsofcapture.Nineofthoseindividualsdiedand,overall,sur- madeextremelydifficultinotherstatessuchasOregonandWashington
vivalwaslowerthanthatofreferenceanimalsthatremainedwithintheir byrestrictionsontheuseofhoundstobringanimalstobay.Nonethe-
originalhomeranges(Ruthetal.1998).Theyconcludedthattranslo- less,carefullyregulatedsporthuntingremainsanimportantrecreational
cationwasmostsuccessfulwhenmountainlionswere12–27months pursuitandisemployedasamethodofsustainingmountainlionpop-
old. Translocation as a management technique for mountain lions is ulations at viable levels in a majority of western states (Logan and
notwidelyimplemented,butmaybecomeincreasinglyimportantasa Sweanor1999).
result of changing attitudes toward predator control (Hancock 1980) Conservationofmountainlionshasalongandvariedhistory,and
andlossorfragmentationofhabitat(NowellandJackson1996). hasbeencouchedlargelyintermsofpoliticalexpediency(Torres1997).
Effortstocontroloreliminatetheseobligatecarnivoresbyunrestricted
Conservation. Conservationofalargecarnivorethatpreysonother harvestandbountysystemswereunsuccessfulthroughoutmuchofthe
largemammals,frequentlyisinvolvedinthekillingofdomesticlive- United States, but their efficacy has been attested to by the absence
stock,andoccasionallyisathreattohumansafetypresentschallenges ofmountainlionsfrommuchoftheirhistoricalrange.Inthoseareas,
thatseemoverwhelmingincontemporarytimes.Theprimarypreyof suchaswesternNorthAmerica,wherelargetractsofessentialhabitat
mountainlionsrequirelargetractsofopenandundevelopedland,both remained in relatively pristine conditions, intensive methods of con-
ofwhicharebecomingincreasinglyuncommon.Mountainlionsthem- trolling mountain lions gave way to regulated sport hunting, with an
selves similarly depend on large tracts of land to meet their needs. emphasisonensuringviablepopulationswiththecapabilityofmain-
Indeed,theconservationandmanagementchallengeforthefuturewill tainingasustainableharvest.Selectiveremovalofindividuallionsthat
betoassurethepresenceofmountainlionsandtheirprey,despitethe have preyed on livestock or domestic animals has been compatible
certain loss of habitat and increases in human numbers and activity throughoutmuchoftherangeofmountainlionsduringtherecentpast.
(Torres1997). Eliminationoflionsinvolvedwithhumansafetyincidentslargelyhas
InwesternNorthAmerica,mountainlionpopulationsgenerally achievedpublicacceptance.
remain healthy (Hornocker 1992), but conservationists will be faced
withthetaskofmaintaininglarge,contiguoustractsofsuitablehabi-
RESEARCHNEEDS
tatthatarelinkedtoothersuchareas.Populationsofmountainlions
mayoccurinametapopulationstructure(Sweanoretal.2000;but,see Mountain lions occur at low densities and are secretive and cryptic;
Ernestetal.2003).Islandsofhabitatsuitableforsupportingpermanent thus,theyareamongthemostdifficultlargemammalstostudy.Fur-
populationssometimesareseparatedbyvastareasofhabitatthatarenot thermore,costsofresearchassociatedwiththestudyoflargecarnivores
permanentlyoccupied,butinsteadprovideopportunitiesformovement aregreat,confoundingeffortstoobtainmeaningfulanswerstodifficult
between habitat islands. Thus, habitat fragmentation is a major con- questions. Although many aspects of the ecology of mountain lions
cern.Beier(1993)andBeieretal.(1995)describedasituationwhere havebeeninvestigated,fewauthorshaveconductedthelong-termin-
mountain lion habitat in the Santa Ana Mountains of southern Cali- vestigationsnecessarytobegintounderstandtherelationshipsamong
forniahadbecomeisolatedfromothersuitableareasandimmigration habitat, prey densities, and the dynamics of the populations of these
likelywasreducedasaresultofhumandevelopment.Immigrationand secretivepredatorsandtheirprey.
emigrationofindividualsarenecessaryforthemaintenanceofgenetic Mountainlionshavebeenimplicatedasfactorsimportantinthe
diversity.Further,opportunitiesforrecolonizationmustbeprovidedin dynamicsofsomepopulationsofmountainsheep(Kamleretal.2002),
theeventofextirpationsfromislandsofsuitablehabitat.Maintaining includingtwopopulationsegments(Wehausen1996;Hayesetal.2000)
adequatespaceformountainlionsandtheirpreyaswellaslinkages listedasendangeredbythefederalgovernment.Impactsoflionpreda-
betweensuchareasclearlyisapressingissuethatmustbeaddressedif tiononmountainsheeppopulationsaremostprobablewheremuledeer
mountainlionsaretoretaintheircurrentstatus. andmountainsheepoccursympatrically(Schaeferetal.2000;Kamler
Depredationsonlivestockandotherdomesticanimalslikelywill etal.2002).Nonetheless,clearlinkagesamongthedynamicsofpop-
increase in frequency, especially as the human population expands ulationsofprimaryprey(muledeer),mountainlions,andsecondary
(Torres1997).Depredationmaybecoincidentwithincreasesinlion prey (mountain sheep) are yet to be established, and warrant serious
populations(Torresetal.1996),butmayalsobeafunctionofdeclines investigation. Such investigations must, however, be conducted over
inpopulationsofprimaryprey(Pierceetal.1999a,Kamleretal.2002). atemporalscaleadequatetoelucidatechangesinhabitatqualityand
Whateverthecause,managementofdepredatingmountainlionswill resultantresponsesamongpreyandpredatorsinsuchsystems(Pierce
remainanimportantissue.Currentmanagementofdepredatingindivid- 1999). As a result, population modeling may become an even more
ualsusuallyinvolveskillingoftheoffendinganimal,astrategyunlikely usefulpredictivetool,(Beier1993)withmorepotentialforapplication
to change in the immediate future. Even in densely populated areas, thancurrentlyrecognized.
sucheventsarerelativelyuncommon(Torres2000),andsuchremovals Thegeneticstructureofpopulationsofmountainlionshasbeen
likelydonotpresentathreattoexistingpopulationsofmountainlions. examined only recently (Walker et al. 2000; Ernest et al. 2003), and
Mountainlionpopulationsapparentlyhaveexpandedduringre- additionalopportunitiestodefinepopulationsfromageneticperspec-
centyears,andtherehasbeenincreasingconcernaboutthepotentialfor tiveareneeded.Furthermore,thespatialstructureoflionpopulations
anincreaseinhumanencounters.Indeed,suchconcernshaveresultedin hasbeenreportedonlyonalocalscale(Germaineetal.2000;Sweanor
publicationofpamphletsandbooks(Torres1997)designedtominimize etal.2000),andlandscape-leveleffortstomoreclearlydefinepopula-
theprobabilityofsuchincidents.Youngmountainlions,dispersingbe- tionsofmountainlionsclearlyarewarranted.Moreover,thepotential
tweenareasofsuitablehabitat,maybemostapttoencounterhumans; metapopulationstructure(Sweanoretal.2000)ofmountainlionswar-
indeed,themajorityofattacksonhumanshaveinvolvedsubadultorap- rantsfurtherinvestigation,andhasimportantimplicationsforthecon-
parentlyunderweightlions(Aune1991;Beier1991).Ashumanpopu- servationoftheselargefelids(Ernestetal.2003).GPStechnologycan
lationsincreaseandexpandintocurrentlyoccupiedlionhabitat,therate be incorporated into telemetry collars for investigations of mountain
atwhichsuchencountersoccurmaybeexpectedtoincrease.Meeting lion movements and habitat selection (Bleich et al. 2000; Anderson
human safety objectives while simultaneously providing habitat for and Lindzey 2003) and can be combined with sophisticated genetic