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Encounter Competition Between Bears and Cougars: Some Ecological

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Encounter Competition Between Bears and Cougars: Some Ecological ENCOUNTERCOMPETITION BETWEEN BEARS AND COUGARS: SOMEECOLOGICAL IMPLICATIONS KERRYM. MURPHY,Hornocker Wildlife Institute, P.O. Box 526, MammothHot Springs,WY 82190, USA, email: kmurph@ gomontana.com GREGORYS. FELZIEN,'Hornocker Wildlife Institute, 3246, UniversityStation, Moscow,ID 83843, WA, USA MAURICEG. HORNOCKER,Hornocker Wildlife Institute, P.O. Box 3246, UniversityStation, Moscow,ID 83843, USA, email: [email protected] TONIK. RUTH,Hornocker Wildlife Institute, 3246, UniversityStation, Moscow,ID 83843, WA, USA, email: RUTH8744@NOVELL. UIDAHO.EDU Abstract: Black bears (Ursus americanus)or grizzly bears (Ursus arctos) visited 8 of 55 cougar-killed (Felis concolor) ungulates in Glacier National Park(GNP), Montana,from 1992 to 1995, and 19 of 58 cougarkills in Yellowstone NationalPark (YNP), Wyoming, from 1990 to 1995. Bears displaced cougars from 4 of 8 carcasses they visited in GNP and 7 of 19 in YNP. Cougar predationprovided an average of 1.9 kg/day (range= 0-6.8 kg/day) of biomass to bears that fed on cougar-killedungulates. This biomass was an importantpercent (up to 113%) of the daily energy needs of bears when comparedto their caloric requirementsreported in the literature. We suggest that ungulate carrionresulting from cougar predationis importantnutritionally to bears in some regions and seasons. Cougars that were displaced from their kills by bears lost an average of 0.64 kg/day of ungulatebiomass, or 17-26% of their daily energy requirements. Biologists modelling or measuringcougar predation rates should be aware that losses to scavengers may be significant. Ursus 10:55-60 Key words: black bear, competition, cougar, Glacier National Park, grizzly bear, predation, Puma concolor, scavenging, Ursus americanus, Ursus arctos, Yellowstone National Park. Encountercompetition is a form of interferencecom- documentthe frequencythat cougars were displacedfrom petitionthat occurs when 1 or more mobile, nonspatially- ungulateprey and calculate the quantityof biomass that attachedindividuals are harmedas they directlypursue a cougars lost to bears. common resource (Schoener 1983). This harmincludes Ourresearch was fundedby G.C. Hixon, M.J.Murdock injuriesresulting from physical interactions,loss of time, CharitableTrust, Montanaof Fish, Wildlife and Parks, or theft of food. For carnivores,the disputed resource Nathaniel P. Reed, the P.W. Busch Family Foundation, may be prey that a subordinatecarnivore has killed and a The NationalFish andWildlife Foundation,The National dominantcarnivore intends to scavenge. Geographic Society, The National Wildlife Federation, Interactionsbetween cougarsand scavengersthat were The R.K. Mellon Foundation,the U.S. Fish and Wildlife presentnear cougar-killed prey have been reported(Boyd Service, and Yellowstone National Park. The Gallatin and O'Gara 1985, Spreadbury 1989, White and Boyd National Forest, GardinerRanger District, and Glacier 1989, Harrison 1990, Green 1994). Koehler and National Park provided logistical support. We thank J. Horocker (1991) found 4 bobcats (Lynx rufus) and 2 Cole, T. Fredrickson,B. Holmes, M. Johnson,J. Jonkel, coyotes (Canislatrans) killed by cougarsnear prey cache J. Murnane, B. O'Gara, T. Parker, S. Relyea, J. sites. Harrison(1990) found that a maternalfemale cou- Tischendorf, and B. Wiesner for their work in the field where gar residing coyote abundance was reduced by or laboratory. B. Chapman and D. Stradley provided humans killed ungulate prey less frequentlythan a ma- aerial support on short notice. P. Munholland played ternalfemale where living coyote numberswere not ma- an important role in designing the cougar predation nipulated. Coyotes displacedcougars from theirkills less studies. D. Pac graciously provided his unpublished frequentlywhere coyote numberswere controlled. data on the weights of mule deer carcasses. We thank Althoughinteractions between cougarsand scavengers agency personnel S. Frye, G. Green, T. Lemke, J. near have been cougar-killedprey documented,the eco- Mack, J. Logan, C. Sime, T. Their, D. Tyers, J. Varley, logical importanceof encountercompetition remains un- and R. Wuertz. We especially thank K. Aune and T. clear. Our was objective to (1) documentthe frequency Moore of the Montana and Wyoming wildlife labora- that black bears and bears used grizzly cougar-killedun- tories, respectively, for their technical support and estimate the biomass gulates, (2) ungulate made avail- guidance. We thank I. Ross and an anonymous per- able to bears as a result of cougar predation, and (3) son for reviewing this manuscript. 1Deceased. 56 Ursus 10:1998 STUDYAREAS 1992-95, in GNP and during spring-autumn,1990-95, in YNP. Using methods reportedby Shaw (1977), cou- GlacierNational Park gars were located 1-3 times daily, between 0600 and Cougar-bearinteractions were documentedconcurrent 2300 hours from the ground or from airplanes,without with studies of cougar-wolf (Canis lupus) relationships disturbing them. All location sites were intensively in a 790-km2area along the North Fork of the Flathead searched by walking the contours of slopes at 1-20 m River in GlacierNational Park (GNP) in northwestMon- intervals to locate carcasses after cougars left. Track- tana. Cougars preyed on elk (Cervus elaphus), white- ing hounds were used to assist searches of locations tailed deer (Odocoileus virginianus), moose (Alces and to trail cougars between location sites. Intact un- alces), and mule deer (Odocoileus hemionus). The val- gulate carcasses were necropsied to distinguish cougar ley was gently rolling terrainsupporting a montanefor- predationfrom that of other carnivoresby looking for est interspersedwith meadows. It was borderedby steep gross pathology or feeding behaviors described in mountains, 1700-3000 m in elevation. Snow accumu- O'Gara (1978), Wade and Bowns (1982), and Alberta lation promptedungulates to winter at elevations <1600 Agriculture(1990). Cougars were monitored on con- m from late November to April. Thereafter,ungulates secutive days until at least 2, but typically 4 ungulates used progressivelyhigher elevations associated with phe- were killed. We attemptedto document all ungulates nological changes in vegetation. killed by cougars during this period, called a predation Black bears and grizzly bears were common. Some sequence. grizzlies were active outside their dens for at least part of the winter, particularlyin the Kintla Lakes area of BearVisits to CougarPrey GNP, where carrionwas relatively abundant. Bear visits to cougar-killedungulates were detected by observing bears or by noting the presence of bear YellowstoneNational Park scats, hair, or tracks near carcasses. Hair samples of Cougarswere studied in the northernportion of Yel- carnivores were identified to genus or species by per- lowstone National Park (YNP), Wyoming, and the con- sonnel at the Wyoming Game and Fish Laboratoryin tiguous ParadiseValley of south-centralMontana. The Laramie,following Moore et al. (1974). We could not winter density of adult-subadultsranged from 2.1 to 3.5 always discernthe species of bear that used cougarprey. per 100 km2 on a 662-km2core area where they spent Displacementof cougarsfrom cougarkills was recorded wintermonths on ungulateranges between 1550-2500 m. as a special case of bear visitation. To identify dis- Elk and mule deer were far more abundantthan moose placement,we comparedthe weight of the ungulatecar- and white-taileddeer. Winteringelk counts rangedfrom cass that remained(after use by the bear) to the amount 9,400 to 19,000 from 1990 to 1995 (Mack and Singer present,on average, after cougars left kills without bear 1992; J. Mack, YNP, unpubl. data). Mule deer counts displacement. When possible, we interpretedtracks of and radiotelem- ranged from 1,600 to 2,500 between 1990 and 1994 cougars and bears visible in the snow (Lemke 1994). Duringlate spring,summer, and autumn, etry data to discern bear displacement. used elevations to 2900 m and on un- cougars up preyed Bears gulates that were dispersedrelative to theirrestricted dis- Biomass Eaten by tributionin winter. Ungulateranges supportedmontane We estimated the wet ungulate biomass eaten by kills in YNP. forest, sagebrushsteppe, and grasslandsteppe. bears (BEB) feeding on cougar only if A minimum of 236 grizzly bears were present in the Biomass was considered available to bears only the 20,000-km2Greater Yellowstone Ecosystem in autumn we detected bears at a cougar kill. BEB was of the 1992 (U.S. Fish and Wildl. Serv. 1993, Eberhardtet al. difference between the live-weight ungulate 1994). Black bears were commonly observed. Bears and the amount consumed by the cougar, multiplied used dens from mid-Novemberto mid-April. by the estimated fractional consumption of carcasses by bears. To approximate the energy content of BEB, we converted BEB to its caloric equivalent by METHODS assuming dry weight was 32.4% of wet weight (Robbins et al. 1974, Robbins 1993) and multiply- DocumentingCougar Predation ing dry weight by its digestible energy of 4.6 kcal/g BEB for each kill was Responses of bears to ungulates killed by (Mealey 1975). cougar and divided the radiocollaredcougars were documented during winter, summed to estimate total BEB by BEARS AND COUGARPREDATION * Murphyet al. 57 length of the predation sequence in days to estimate on a small numberof observations,(2) coyotes, avifauna, daily BEB. decomposers,tissue autolysisand decay, andwater evapo- Weightsof UngulatePrey.-The weights of ungulates ration reduced carrionbiomass, particularlyif bears in- killed by cougars were estimatedusing
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