Survival and Mortality of Translocated Woodland Caribou Author(S): Bradley B

Home , Selkirk Mountains

Survival and Mortality of Translocated Woodland Caribou Author(s): Bradley B. Compton, Peter Zager and Gregg Servheen Reviewed work(s): Source: Wildlife Society Bulletin, Vol. 23, No. 3 (Autumn, 1995), pp. 490-496 Published by: Allen Press Stable URL: http://www.jstor.org/stable/3782959 . Accessed: 07/11/2012 15:12

Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp

. JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected].

Allen Press is collaborating with JSTOR to digitize, preserve and extend access to Wildlife Society Bulletin.

http://www.jstor.org Caribou survival

Survival and mortality of translocated woodland caribou

Bradley B. Compton, Peter Zager, and Gregg Servheen

AbstractWe monitoredsurvival of 60 woodland caribou (Rangifertarandus) translocatedfrom BritishColumbia to the SelkirkMountains of northernIdaho betweenMarch 1987 and February1992. This translocationwas to assistin recoveryof the endangeredSelkirk population. For all translocatedcaribou combined,estimated annual survivalrates rangedfrom 0.65-0.94 and wereconsistent with declining established populations. No differences(P > 0.10) in survivalwere foundbetween male and femalecaribou or be- tweenmountain and northernecotypes. Causes ofdeath included unknown (n = 14), predation(n = 7), other(n = 4), and human-caused(n = 2), withsummer accounting for the greatestproportion (53o/o). Emigrationof 7 mountainecotype animals also was a loss to the population. Mountainlions (Fe/is concolor) caused mostconfirmed predator kills. Seasonalpattern of mortality was consistentwith established populations where predation was identifiedas a significantfactor.

Keywords caribou,mortality, Rangifer tarandus, survival, translocation

Historically,woodland caribou(Rangifer tarandus Washington,and southernBritish Columbia, Canada caribou) rangedover much of the northerntier of the (Scottand Servheen1985). Thispopulation was listed UnitedStates (U.S. Fishand Wildl.Serv. [USFWS] 1985). as endangeredin 1984 underthe EndangeredSpecies Bythe early 1980's, their U.S. distributionhad been re- Act. Human-causedmortality, habitat loss, and forest duced to a smallherd of 25-30 animalsinhabiting the fragmentationwere believedto be limitingthe Selkirk SelkirkMountains of northernIdaho, northeastern population(USFWS 1985). The managementand re- coveryplan (USFWS1985) identified 4 objectivesfor the Selkirkpopula- tion: (1) controlpoaching; (2) mini- mizecaribou-vehicle collisions along BritishColumbia Highway 3; (3) im- prove habitatquality; and (4) aug- .'4 mentthe population. Whiletranslocation is potentially an importanttool forspecies conservation(Griffith et al. 1989), littleinfor- mationhas been reportedon post-release survivalrates of caribou(Klein 1968,Leader-Williams 1988, Bergerud and Mercer1989). Betterinformation and understandingof survivalrates is neededto evaluatethe role of translo- cationfor conserving the Selkirkcari- bou and otherpopulations.

All authorsare withthe Idaho Departmentof Fishand Game. BradleyB. Compton: 1345 BartonRoad, Pocatello,ID 83204, USA; Peter Zager: 1540 WarnerAvenue, Lewiston, ID 83501, USA; GreggServheen: 1540 WarnerAvenue, Lewiston,ID 83501, USA. All photos fromIdaho Fishand Game.

WildlifeSociety Bulletin 1995, 23(3):490-496 Peer refereed Caribousurvival * Comptonet al. 491

Our objectivewas to monitorthe survivalof cari- northernIdaho, and southernBritish Columbia (see bou moved fromBritish Columbia (B.C.) to unoccu- map). Approximately2,700 km2 (47%o)of this area pied habitatin the SelkirkMountains of Idaho (U.S. lies in B.C. and 3,000 km2 (53%) are in the U.S., in- For. Serv. [USFS] 1985). We reportsurvival rates and cludingthe Salmo-PriestWilderness, portions of the mortalitycauses fortranslocated caribou from March Colvilleand Idaho PanhandleNational forests, Idaho 1987-February1992. Departmentof Lands holdings,and scatteredprivate parcels. Physiographyis characterized by short, steep-sloped drainages. Elevations range from Study area 540-2,375 m. The Pacificmaritime-continental cli- Capture sites mate is characterizedby long wintersand shortsum- All capturedcaribou came fromBritish Columbia. merswith an averageannual precipitation of ?250 cm Caribou were captured duringthe wintersof 1987 and wintersnowfall generally >700 cm at higherele- and 1988 fromthe Ilgachuz Mountainsnear Anahim vations(Krajina 1967). Vegetationis dominatedby Lake and the northern Selkirk Mountains near Engelmannspruce-subalpine firand western red- Revelstoke.During the winter of 1990, caribouwere cedar-westernHemlock foresttypes (Cooper et al. capturedfrom the Cariboo and Monasheemountains 1987). Caribouin thisecosystem, characterized by ar- near Blue River. AnahimLake is approximately580 boreal lichens and heavy snowfall,are classifiedas km northwest of the Selkirks. Blue River and mountainecotype". Revelstokeare approximately420 and 285 kmnorth- northwest,respectively. The Ilgachuz Mountains are located within the Methods InteriorPlateau region of west-central B.C. witheleva- Fourteencaribou were capturedduring March by tions rangingfrom 1,200-2,400 m (Holland 1976). drive-netting(Beasom et al. 1980) and 49 by net-gun- Biotic zones are dominatedby the sub-borealpine ning (Barrettet al. 1982). Captured caribou were (lodgepole pine, Pinus contorta)-spruce (white manually restrained, blindfolded, sedated with spruce,Picea glauca and Engelmannspruce, P. engel- 40-100 mg xylazinehydrochloride, and flown to a mannfi), Engelmannspruce-subalpine fir (Abies la- holdingpen. At the holdingpen, each caribouwas siocarpa), and alpinetundra zones. Climateis charac- fittedwith a mortality-sensingradio collar (Telonics, terizedby cool, short,dry summers; winters are cold, Inc., Mesa, Ariz.) and numbered plastic ear tags long,and dry. Precipitationaverages 42 cm annually (AllflexInt., Ltd., New Zealand). All animalswere withwinter snowfall averaging 50 cm. Cariboufrom classifiedas eithercalves (<1 year)or adults(?1 year). the Ilgachuz Mountainsprimarily rely on terrestrial lichens (Cladina spp., Cladonia spp., Stereocaulon spp.,Peltigera spp.) duringwinter and are considered "northernecotype" (Stevenson and Hatler1985). The northern Selkirk, Cariboo, and Monashee mountainslie withinthe InteriorWet Belt regionof southeasternB.C. with elevations from460-3,050 NELSONi meters(Krajina 1967). Engelmannspruce-subalpine fir,western redcedar (Thuja pl/cata)-westernhem- KOOTENAY lock (Tsuga heterophylla),and alpine tundraare the dominantbiotic zones. The wet continentalclimate Highway 3 1 N is characterizedby moist,warm summersand cold winterswith heavy snowfall. Annual precipitation BRITISH COLUMBIA ] rangesfrom 56-280 cm with snowfallgenerally ex- WASH. IDAHO km ceeding 700 cm. Cariboufrom this region primarily . Q ~ ~~~25 use arboreal lichens (Alectoria spp., Bryoria spp.) Ball duringthe winterand are referredto as "mountain l Creek ii ecotype"(Stevenson and Hatler1985). o s \ ~BONNERS ~~~~~FERRY Release site Occupied and potentialwoodland caribou habitat was approximately 5,700 km2 in the Selkirk SelkirkMountains woodland caribou habitat in the U.S. and Mountains, located in northeasternWashington, Canada. 492 WildlifeSociety Bulletin 1995, 23(3):490-496

-j W - 7 - - T, the population. Deaths were attributedto predation when carcasseswere in good condition,had signsof predatorfeeding (e.g. caching,tissue removal),der- mal puncturewounds withhemorrhaging, and exhib- ited no grossly identifiablepredisposing factors. Unknowncauses of mortalitywere generallyassoci- ated with extensively scavenged carcasses. Ad- ditionally,deaths were attributedto unknowncause when necropsycould not identifygross morphological or physiologicalimpairments and marrowfat content was <15%. Deaths were attributedto other causes when marrowfat contentwas <15% (Davis and Valkenburg1985, Davis et al. 1987) or necropsy indicatedgross naturally occurring morphological or physiologicalimpairments. Predation and human- caused mortalitywas not ruledout in manyof the unknown cases. Likewise,other causes of death were not ruled out, although malnourishment(as evi- denced bybone marrowfat content) was notbelieved to be a major contributor(Hanks 1981, Davis and Valkenburg1985). Annual (March throughFebruary) survival rates and 5-yearsurvival functions were estimatedwith the Kaplan-Meierstaggered entry design (Pollock et al. Cariboubeing prepared for relocation from British Columbia, 1989). Caribousurvival data were censored(Pollock to U.S.A. Canada, Idaho, et al. 1989, Whiteand Garrott1990) forloss-failure of the transmitter.Differences in survivalfunctions Blood sampleswere takenfor disease titersand tu- were evaluatedusing chi-squareanalysis (Pollock et berculinwas administeredfor tuberculosis testing. al. 1989). Significancewas determinedat P < 0.10. Additionally,caribou were administered1,000,000 units (u) VitaminA, 150,000 u VitaminD, 400 u VitaminE, 2.5 ml VitaminB complex, 6.76 mg sele- Results nium,590,000 u short-actingantibiotics, 1,350,000 u Sixtycaribou (15 adultmales, 1 male calf,42 adult long-actingantibiotics, and 21.1I mg Ivermectin. Fecal females,2 femalecalves) were translocated,includ- samples were collected opportunisticallyto test for ing 24 in 1987, 24 in 1988, and 12 in 1990 (Table 1). presence of meningealworm (Parelaphostrongylus Three caribou died duringcapture and transportop- tenuis). Afterprocessing, caribou were held in 2 cir- erations. No evidence was detectedof tuberculosis, cular holding pens with snow or ice and arboreal brucellosis,or meningealworm. Survivaldata were lichensprovided ad libitum. After72 hours,caribou censored for 12 animalswith transmitterloss or fail- were examinedfor reaction to tuberculinand loaded ure and 1 death caused by a largeice-ball forming on intodarkened horse trailers for transport. the radio collar. Caribouwere released in the Ball Creek drainage, BoundaryCounty, Idaho. Servheen(1 987, 1988) and Table 1. Woodland caribou moved fromBritish Columbia to Compton et al. (1990) provided further details on Idaho, 1987-1990. caribou capture, holding,and transport. Afterre- lease, caribouwere monitoredweekly (weather per- Males Females mitting)from fixed-wing aircraft. Mortalitieswere located on theground and necrop- Capturesite Year Total Adult Calf Adult Calf sied to determinecause of death. Major leg bones AnahimLake 1987 12 2 10 were collectedfor marrow fat content (Nieland 1970) 1988 14 4 10 and lower jaws forage determination(Davis 1978). Revelstoke 1987 12 2 9 1 We classifiedcauses of mortalityas: predation,un- 1988 10 3 1 5 1 known, human-caused,and other. Emigrationfrom Blue River 1990 12 4 8 the ecosystemwas considereda mortalitysource for Total 60 15 1 42 2 Caribousurvival v Comptonet al. 493

Survivalwas calculatedusing 26 deathsand 7 emi- Table 3. Sources of loss among translocatedcaribou by season,' grations. All 7 emigrationswere mountainecotype, March 1987-February1992, in the SelkirkMountains of Idaho. and 6 of these emigratedwithin 1 year post-release. Cause Winter Spring Summer Fall Total Fourteencaribou with active radio collarsremained alive at the end of the reportingperiod. For all cari- Unknown 3 2 9 0 14 bou combined,the 5-yearweighted average survival Predation 2 0 5 0 7 was 0.74 (SE = 0.10), rangingfrom 0.65-0.94 annu- Emigration 1 1 4 1 7 allywith no trendevident (Table 2). Males and fe- Other 1 2 0 1 4 Human-caused 1 0 0 1 2 males had 5-yearweighted average annual survival Total 8 5 18 3 34 ratesof 0.76 (SE = 0.20) and 0.73 (SE = 0.13), which did not differ(X2 = 0.00, 1 df,P = 0.99). Annualsur- a Winter:1 Nov-15 May; Spring:16 May-15 July;Summer: vival rates for mountain caribou ranged from 16 July- 15 Sept; Fall: 16 Sept-31 Oct. 0.56-0.91 with a 5-yearweighted annual average of 0.74 (SE = 0.13). Annualsurvival for northern caribou ranged from0.58-1.00 with a 5-yearweighted were not available for the remaining3. Of the 4 averageof 0.73 (SE = 0.12). Survivalrates did not dif- deathsattributed to othercauses; 2 revealedevidence fer between mountainand northerncaribou (X2 = of malnourishment,1 showed severe parasitism 0.00, 1 df,P = 0.97). (Sarcocystissp.) and emaciation,and 1 had birthcom- Excluding emigrations,27 caribou died during plications. Predationwas ascribed to 7 of the 27 March 1987-February1992 (Table 3). Of these 27 deaths,including 5 to mountainlion (Felis concolor) deaths,8 (30%) were in the 0-3 age class, 14 (52%) in and 2 to bear (Ursus spp.). Predationwas ruledout as the 4-9 age class, 1 (4%) in the 10+ age class, and 4 a possible cause of death in 3 of 14 unknowncases. (15%) were of unknownage Human-causedmortalities included 1 illegalkill and Summeraccounted for 53% (14 of 27) of all deaths the previouslynoted collar with an ice-ball. (Table 3). Fourteenof 27 (53%) deathswere attrib- uted to unknown causes because scavengers de- stroyedevidence. Malnourishmentwas notevident in Discussion 11 of these 14 deaths(Franzmann and Arneson1976, Althoughseveral translocationsof caribou have Davis and Valkenburg1985). Bone marrowsamples been conducted (Klein 1968, Leader-Williams1988, Bergerudand Mercer 1989, M. McCollough,Maine caribouproject, 1986-1990, FinalRep., Me. Caribou Table 2. Annual(March through February) survival rates of wood- Proj., Portland,1991), we report survivalrates of land caribou translocatedfrom March 1987 throughFebruary 1992, SelkirkMountains, Idaho. translocatedwoodland caribou for the firsttime. Generally,survival rates <0.85 for establishedcari- Class n S 90% Cl bou are characteristic of declining populations (Bergerud1978, 1980, 1983). Our estimatesof an- All caribou 1987-1988 24 0.65 0.48-0.82 nual survivalfor all translocatedcaribou combined 1988-1989 38 0.82 0.71 -0.92 were below thisthreshold value 4 of5 years. Withan 1989-1990 29 0.66 0.50-0.82 approximatemaximum intrinsic rate of increase of 1990-1991 28 0.67 0.52-0.82 0.30 (Bergerud 1980), adult survivalmust exceed 1991 -1992 17 0.94 0.84-1.00 0.70 forcaribou populations to sustainthemselves. Ecotype Mountain Unfortunately,we do nothave reliablerecruitment 1987-1988 12 0.56 0.31 -0.80 informationfrom 1987-1990 to compare with our 1988-1989 16 0.87 0.73-1.00 survivalestimates. However, data collected on the 1989-1990 13 0.75 0.53-0.97 residentSelkirk population from1983-1985 (Scott 1990-1991 20 0.63 0.45-0.81 and Servheen 1985) suggested annual recruitment 1991-1992 11 0.91 0.77-1.00 Northern rates between 0.12 and 0.18. Additionally,recruit- 1987-1988 12 0.73 0.51 -0.95 mentestimates for 1991 and 1992 were 0.14 and 0.06 1988-1989 22 0.77 0.62-0.92 (B. B. Compton,P. Zager, and J. R. Skalski,Selkirk 1989-1990 16 0.58 0.36-0.80 Mountains woodland caribou census technique- 1990-1991 8 0.75 0.50-1.00 methodologyand recommendations. Idaho Dep. 1991 -1992 6 1.00 Fish and Game, Boise, 1992). Withthis range of re- a n indicatesthe numberof individualsknown to be alive at the cruitment,it is unlikelythat production offset mortal- beginningof the annual period. ity,further suggesting a decliningpopulation. 494 WildlifeSociety Bulletin 1995, 23(3):490-496

Survivalof translocatedanimals may be less than woodland caribou. We found that mountainlion that of residentanimals, as was found with white- are also importantpredators. tailed deer (Odocoileus virginianus) in Illinois Bergerudand Elliot (1986) and Seip (1992) sug- (Jones and Witham1990) and black-taileddeer (0. gested that caribou experience higher predation hemionus columbianus) in California(O'Bryan and rates in multiple prey-predator systems. McCullough 1985). Six radio-collared resident Specifically,they indicated that moose (Alces alces) Selkirkcaribou (Scott and Servheen 1985) had a 5- support higherwolf numberswhich subsequently year (March 1983-February1988) annual weighted increases predationon caribou. The Selkirkstudy average survivalof 0.94 (SE = 0.08). However, the area is a multiple prey-predatorsystem. White- survivalfunction did not differfrom that of translo- taileddeer, elk (Cervus elaphus), moose, and moun- cated caribou (x2 = 2.70, 1 df,P = 0.11). Limited tainlion populationshave apparentlyincreased over sample size maynot have allowed us to detect a dif- recent historicallevels (Idaho Fish and Game, un- ference. publ. data). Black bear (U. americanus) also are Mountainecotype caribou exhibitdifferent habi- abundant within the Selkirkecosystem (Rohlman tat use patterns,seasonal migrations,and winter 1989). diet selections than northern ecotype animals (Stevenson and Hatler 1985, Servheen and Lyon 1989). Our resultsfor mountain and northerneco- Conservation implications typeanimals suggest there was no augmentationad- Success of the Selkirkaugmentation project can vantageto usingeither ecotype. Over the 5-yearpe- be evaluated in 3 phases. Phase I is whether riod, 50% of the mountainecotype died compared translocatedcaribou successfullyoccupied and re- to 62% of the northernecotype; however, emigra- produced within the ecosystem. Phase II is tion of mountain ecotype made up for perceived whethera short-termself-sustaining population re- survivaladvantages. sulted. Phase III is the ultimate recovery of the Predominantlysummer mortalityhas been re- population. Our resultsindicate that phase I was ported for woodland caribou populations in Wells successful. However, phase II may not occur be- Gray Provincial Park (Seip 1990), Quesnel cause of low survivalrates. We recommend that Highlands (Seip 1992), and west-centralAlberta monitoringof the remainingradio-collared animals, (Edmonds 1988). In contrast,Simpson and Woods theiroffspring, and the entirepopulation continue, (1987) in the Columbia Riverdrainage, Farnell and emphasizing survival,recruitment, and causes of McDonald (1988) in the Yukon, and Fuller and mortality. Keith (1981) in northeastern Alberta reported Before furtheraugmentations are implemented, highermortality during other seasons. Studies re- we recommendthat a thoroughhabitat evaluation be portingpredominantly summer mortality indicated conducted. Not only should structuralcomponents primarycauses of death were human-causedor pre- of habitatbe evaluated,but also the functionalas- dation. Our data, with a large numberof unknown pects. These functionalaspects include but are not causes of death, fail to clearly identifya primary limited to the relationships among habitat- cause of mortality.However, we believe humans prey-predatorand potentialrange overlap of caribou were not a significant factor. Likewise, and predators. Parelaphostrongylus tenuis was not a factor as Foreytand Compton (1991) found no incidence of Acknowledgments. We thank M. Gratson, D. meningealworm in a sympatricSelkirk population Leptich,W. Wakkinen,K. Whitten,and 3 anonymous of white-taileddeer. Additionally,malnourishment reviewersfor providing critical improvements to the is unlikelyduring summer. Predationaccounted for manuscript.K. Burns,J. Gallagher,and G. Beck pro- at least 26% of translocatedcaribou deaths. We be- vided manysafe hours of flying. C. Warrenand L. lieve this is a conservativeestimate because preda- Allen-Johnsonassisted with data collection. Finally, tion was most likelyresponsible in manyof the un- we thankT. Reinecker,L. Kuck,and W. Melquistfor known cases. Our seasonal patternof mortalityis theirsupport. This studyis a contributionof Section most consistentwith otherwork identifyingpreda- 6, Endangered Species Act, Threatened and tion as a significantcause of mortality(Bergerud Endangered Species Project E-7, USFWS; Idaho 1974, 1980, 1983; Fullerand Keith 1981; Bergerud Departmentof Fish and Game, U.S. Forest Service and Elliot 1986; Farnell and McDonald 1988; Seip British Columbia Ministry of Environment, 1990, 1992). Graywolves (Canis lupus) and bears WashingtonDepartment of Wildlife, and Washington have long been consideredthe primarypredators of StateUniversity cooperating. Caribousurvival d Comptonet al. 495

HANKS, J. 1981. Characterizationof populationcondition. Pages Literaturecited 47-73 in C. W. Fowler and T. D. Smith,eds. Dynamics of BARRETT,M. W., J.W. NOLAN,AND L. D. Roy. 1982. Evaluationof a large mammal populations. John Wiley and Sons, New hand held net-gunto capturelarge mammals. Wildl. Soc. Bull. York,N.Y. 10:108-114. HOLLAND, S. 1976. Landformsof BritishColumbia: a physio- BEASOM,S. L.,W. EvANs,AND L. TEMPLE. 1980. The drivenet for cap- graphicoutline. BritishColumbia Dep. Mines and Petroleum turingwestern big game. J.Wildl. Manage. 44:478-480. Resour.Bull. 48. 138pp. BERGERUD, A. T. 1974. Decline of caribou in NorthAmerica fol- JONES, J.M., AND J.H. WITHAM. 1990. Post-translocationsurvival lowingsettlement. J. Wildl. Manage. 38:757-770. and movementsof metropolitanwhite-tailed deer. Wildl.Soc. BERGERUD, A. T. 1978. Caribou. Pages 83-101 inJ. L. Schmidtand Bull. 18:434-441. D. L. Gilbert,eds. Big game of NorthAmerica: ecology and KLEIN, D. R. 1968. The introduction,increase, and crash of rein- management.Stackpole Books, Harrisburg, Pa. deer on St. MatthewIsland. J.Wildl. Manage. 32:350-367. BERGERUD, A. T. 1980. A reviewof the population dynamics of cari- KRAJiNA,V. J. 1967. Biogeoclimaticzones and classificationof bou and wild reindeer in North America. Proc. Int. BritishColumbia. Ecol. WesternNorth Am. 1:1-17. Reindeer/CaribouSymp. 2:556-581. LEADER-WILLIAMS,N. 1988. Reindeeron SouthGeorgia: the ecology BERGERUD, A. T. 1983. The naturalpopulation control of caribou. of an introducedpopulation. CambridgeUniv. Press, New Pages 14-61 in F. L. Bunnell,D. S. Eastman,andJ. M. Peek,eds. York,N.Y. 319pp. Proc. of symposium: Naturalregulation of wildlifepopula- NIELAND, K. A. 1970. Weightof driedmarrow as indicatorof fat in tions. For.,Wildl., and RangeExp., Stn.,Univ. Idaho, Moscow. cariboufemurs. J. Wildl. Manage. 34:904-907. BERGERUD, A. T., AND J. P. ELLIOT. 1986. Dynamicsof caribou and O'BRYAN, M. K., AND D. R. MCCULLOUGH. 1985. Survivalof black- wolves in northern British Columbia. Can. J. Zool. taileddeer followingrelocation in California.J. Wildl. Manage. 64:1,515-1,529. 49:115-119. BERGERUD, A. T., AND W. E. MERCER. 1989. Caribouintroductions in POLLOCK, K. H., S. R. WINTERSTEIN, C. M. BUNCK, AND P. D. CURTIS. easternNorth America. Wildl.Soc. Bull. 17:111-120. 1989. Survivalanalysis in telemetrystudies: the staggered COMPTON, B. B., P. ZAGER, AND L. ALLEN-JOHNSON. 1990. Selkirk entrydesign. J.Wildl. Manage. 53:7-15. Mountainscaribou transplant, October 1989-September1990. ROHLMAN, J.A. 1989. Black bear ecology near PriestLake, Idaho. Annu. Rep., Threatenedand EndangeredSpecies Proj. E-7-1. M.S. Thesis,Univ. Idaho, Moscow. 76pp. Idaho Dep. Fishand Game,Boise. 31pp. SCOTT,M. D., AND G. L. SERVHEEN. 1985. Caribouecology. Pittman- COOPER, S. V., D. E. NEIMAN, R. S. STEELE, AND D. W. ROBERTS. 1987. RobertsonJob Comp. Rep. Proj. W-160-R-12.Idaho Dep. Fish Foresthabitat types of northern Idaho: a second approximation. and Game,Boise. 136pp. U.S. Dep. Agric.For. Serv.Gen. Tech. Rep. INT-236. 135pp. SEIP, D. R. 1990. Ecology of woodland caribou in Wells Gray DAvIS,J. L. 1978. Age structureof caribou populationsin arctic ProvincialPark. Wildl. Bull. B-68. BritishColumbia Ministr. Alaska. Fed. AidWildl. Restor. Prog. Final Rep. Proj.W-17-9, W- Environ.,Victoria. 43pp. 17-10. AlaskaDep. Fish and Game,Juneau. 9pp. SEop,D. R. 1992. Habitatuse and populationstatus of woodland DAVIS,J. L., AND P. VALKENBURG. 1985. Qualitativeand quantitativeas- caribou in the Quesnel Highlands,British Columbia. Wildl. pects of naturalmortality of the WesternArctic caribou herd. Bull. B-71. BritishColumbia Ministr. Environ., Victoria. 50pp. Fed. AidWildl. Restor. Prog. Final Rep. Proj.W-17-1 1, W-21-2, W- SERVHEEN, G. L. 1987. Selkirk Mountain caribou transplant, 22-1,W-22-2, W-22-3. Alaska Dep. Fishand Game,Juneau.71pp. October 1986-September1987. Annu.Rep., Threatenedand DAvis,J. L., P. VALKENBURG,AND D. J. REID. 1987. Correlationsand EndangeredSpecies Proj. Idaho Dep. Fish and Game, Boise. depletionpatterns of marrowfat in caribou bones. J. Wildl. 21pp. Manage. 51:365-371. SERVHEEN,G. L. 1988. SelkirkMountains caribou transplant, October EDMONDS, J. E. 1988. Populationstatus, distribution, and move- 1987-September1988. Ann.Rep., Threatenedand Endangered ments of woodland caribou in west centralAlberta. Can. J. Species Proj. Idaho Dep. Fishand Game,Boise. 40pp. Zool. 66:817-826. SERVHEEN,G. L.,AND L. J.LYON. 1989. Habitatuse bywoodland cari- FARNELL,R., AND J. McDONALD. 1988. The influenceof wolfpreda- bou in the SelkirkMountains. J. Wildl. Manage. 53:230-237. tion on caribou mortalityin Yukon's Finlaysoncaribou herd. SIMPSON, K., AND G. P. WOODS. 1987. Movementsand habitatsof Pages 52-72 in R. D. CameronandJ. L. Davis, eds. Proc. North caribouin the mountainsof southernBritish Columbia. Wildl. Am. CaribouWorkshop 3:52-72. Bull. B-57. BritishColumbia Ministr. Environ., Victoria. 36pp. FOREYT,W. J.,AND B. B. COMPTON. 1991. Surveyfor meningeal STEVENSON, S. K., AND D. F. HATLER. 1985. Woodland caribou and worm (Parelaphostrongylustenuis) and ear mites in white- their habitat in southern BritishColumbia. Vol. 1. Land taileddeer fromnorthern Idaho. J.Wildl. Dis. 27:716-718. Manage. Rep. 23. BritishColumbia Ministr. Forests, Victoria. FRANZMANN,A. W., AND P. D. ARNESON. 1976. Marrowfat in Alaskan 355pp. moose femursin relationto mortalityfactors. J. Wildl. Manage. U.S. FOREST SERVICE. 1985. SelkirkMountains caribou herd aug- 40:336-339. mentation:a cooperativeinteragency plan. Idaho Panhandle FULLER,T. K., AND L. B. KEITH. 1981. Woodlandcaribou population Natl.Forests, Coeur d'Alene. 62pp. dynamics in northeastern Alberta. J. Wildl. Manage. U.S. FISH AND WILDLIFE SERVICE. 1985. SelkirkMountain caribou 45:197-213. managementplan. U.S. Fish Wildl. Serv., Portland,Oreg. GRIFFITH, B., J. M. SCOTT, J. W. CARPENTER, AND C. REED. 1989. 118pp. Translocationas a species conservationtool: statusand strat- WHITE, G. C., AND R. A. GARROTT.1990. Analysisof wildliferadio- egy. Science 245:477-480. trackingdata. AcademicPress Inc., San Diego, Calif. 383pp. 496 WildlifeSociety Bulletin 1995, 23(3):490-496

Bradley(Brad) B. Comp- is currentlyevaluating white-tailed deer buck mortality in northern ton(photo) is a Regional Idahoand supervising several other big game and upland bird inves- Wildlife Biologist with tigations.Gregg Servheen is a RegionalWildlife Biologist with the

'r' IX - ' MS:5 _ _ the Idaho Departmentof IdahoDepartment ofFish and Game. He receiveda B.S.in Wildlife Fish and Game. He re- Biologyat theUniversity ofMassachusetts and an M.S. inFisheries ceived a B.S. in Biology andWildlife Sciences at Texas A&M University. His professional in- and an M.S. in Fish and terestsinclude big gameand nongamespecies and developing Wildlife Management ecosystemmanagement through improved management effective- from Montana State ness. University. His professional interestsinclude populationdynamics and managementof cervids. Peter (Pete) Zager is a PrincipalWildlife Re- search Biologistwith the Idaho Departmentof Fish and Game. He completeda B.S. and an M.A. in Biologyat Western MichiganUniversity and a Ph.D. at the Universityof Montana. He AssociateEditor. Feldhamer.