Skip to main content Accessibility help

We use cookies to distinguish you from other users and to provide you with a better experience on our websites. Close this message to accept cookies or find out how to manage your cookie settings.

Close cookie message
×
Hostname: page-component-546b4f848f-w58md Total loading time: 0 Render date: 2023-06-03T04:45:29.627Z Has data issue: false Feature Flags: { "useRatesEcommerce": true } hasContentIssue false

Chapter 13 - Brown Bear (Ursus arctos; North America)

from Part II - Species Accounts

Published online by Cambridge University Press:  16 November 2020

By
Mark A. Haroldson ,
Melanie Clapham ,
Cecily C. Costello ,
Kerry A. Gunther ,
Katherine C. Kendall ,
Sterling D. Miller ,
Karine E. Pigeon ,
Michael F. Proctor ,
Karyn D. Rode  and
Christopher Servheen
...Show all authors
Edited by
Vincenzo Penteriani  and
Mario Melletti
Vincenzo Penteriani
Affiliation:
Spanish Council of Scientific Research (CSIC)
Mario Melletti
Affiliation:
WPSG (Wild Pig Specialist Group) IUCN SSC
Get access

Summary

This chapter comprises the following sections: names, taxonomy, subspecies and distribution, descriptive notes, habitat, movements and home range, activity patterns, feeding ecology, reproduction and growth, behavior, parasites and diseases, status in the wild, and status in captivity.

Keywords

brown bearUrsus arctostaxonomydistributionhabitatbehaviorconservationstatusNorth America
Type
Chapter
Information
Bears of the World
Ecology, Conservation and Management
 , pp. 162 - 195
Publisher: Cambridge University Press
Print publication year: 2020

Access options

Get access to the full version of this content by using one of the access options below. (Log in options will check for institutional or personal access. Content may require purchase if you do not have access.)

References

Alberta Environment and Parks. (2016). Alberta grizzly bear (Ursus arctos) draft recovery plan. Alberta Species at Risk Recovery Plan No. 38. Edmonton: Alberta Environment and Parks.Google Scholar
Ashrafzadeh, M. R., Kaboli, M. & Naghavi, M. R. (2016). Mitochondrial DNA analysis of Iranian brown bears (Ursus arctos) reveals new phylogeographic lineage. Mammalian Biology 81: 1–9.CrossRefGoogle Scholar
Aune, K. E. & Kasworm, W. (1989). East front grizzly bear study. Final report. Helena, MT: Montana Fish, Wildlife and Parks.CrossRefGoogle Scholar
Aune, K. E., Mace, R. D. & Carney, D. W. (1994). The reproductive biology of female grizzly bears in the Northern Continental Divide Ecosystem with supplemental data from the Yellowstone Ecosystem. International Conference on Bear Research and Management 9: 451–458.Google Scholar
Ballard, W. B., Spraker, T. H. & Taylor, K. P. (1981). Causes of neonatal moose calf mortality in south-central Alaska. Journal of Wildlife Management 45: 335–342.Google Scholar
Ballard, W. B., Ayres, L. A., Roney, K. E., Reed, D. J. & Fancy, S. G. (1991). Demography of Noatak grizzly bears in relation to human exploitation and mining development. Federal Aid in Wildlife Restoration Research Final Report for Grants W-22-5, W-22-6, W-23-1, W-23-2, W-23-3 Study 4.20. Juneau, AK: Alaska Department of Fish and Game, Division of Wildlife Conservation.Google Scholar
Ballard, W. B., Ayres, L. A., Reed, D. J., Fancy, S. G. & Roney, K. E. (1993). Demography of grizzly bears in relation to hunting and mining development in Northwestern Alaska. Scientific Monograph Series NPS/NRARO/NRSM-93/23. Denver, CO: National Park Service.Google Scholar
Banci, V. (1991). The status of the grizzly bear in Canada in 1990. Ottawa: Committee on the Status of Endangered Wildlife in Canada.Google Scholar
Barker, O. E., Derocher, A. E. & Edwards, M. A. (2015). Use of Arctic ground squirrels (Urocitellus parryii) by brown bears (Ursus arctos). Polar Biology 38: 369–379.CrossRefGoogle Scholar
Barnes, V. G. Jr. (1990). The influence of salmon availability on movements and range of brown bears on southwest Kodiak Island. International Conference on Bear Research and Management 8: 305–313.Google Scholar
Barnes, V. G. & Van Daele, L. J. (2006). Productivity of female brown bears on Kodiak Island, Alaska. Job completion report for Challenge Cost Share Project. Kodiak, AK: Kodiak National Wildlife Refuge, US Fish and Wildlife Service.Google Scholar
British Columbia Ministry of Forests, Lands, and Natural Resource Operations. (2012). 2012 Grizzly bear population estimate for British Columbia. Victoria: British Columbia Ministry of Forests, Lands, and Natural Resource Operations.Google Scholar
Belant, J. L., Griffith, B., Zhang, Y., Follmann, E. H. & Adams, L. G. (2010). Population-level resource selection by sympatric brown and American black bears in Alaska. Polar Biology 33: 31–40.CrossRefGoogle Scholar
Bellemain, E., Swenson, J. E. & Taberlet, P. (2006). Mating strategies in relation to sexually selected infanticide in a non-social carnivore: the brown bear. Ethology 112: 238–246.CrossRefGoogle Scholar
Ben-David, M., Titus, K. & Beier, L. R. (2004) Consumption of salmon by Alaska brown bears: a trade-off between nutritional requirement and the risk of infanticide? Oecologia 138: 465–474.CrossRefGoogle ScholarPubMed
Bjornlie, D. D., van Manen, F. T., Ebinger, M. R., et al. (2014). Whitebark pine, population density, and home-range size of grizzly bears in the Greater Yellowstone Ecosystem. PloS ONE 9: e88160.CrossRefGoogle ScholarPubMed
Blanchard, B. M. (1987). Size and growth patterns of the Yellowstone grizzly bear. International Conference for Bear Research and Management 7: 99–107.Google Scholar
Blanchard, B. M. & Knight, R. R. (1980). Status of grizzly bears in the Yellowstone system. Transactions of the North American Wildlife and Natural Resources Conference 45: 263–267.Google Scholar
Blanchard, B. M. & Knight, R. R. (1991). Movements of Yellowstone grizzly bears. Biological Conservation 58: 41–67.CrossRefGoogle Scholar
Blanchard, B. M. & Knight, R. R. (1995). Biological consequences of relocating grizzly bears in the Yellowstone Ecosystem. Journal of Wildlife Management 59: 560–565.CrossRefGoogle Scholar
Boertje, R. D., Gasaway, W. C., Grangaard, D. V. & Kellyhouse, D. G. (1987). Predation on moose and caribou by radio-collared grizzly bears in east central Alaska. Canadian Journal of Zoology 66: 2492–2499.CrossRefGoogle Scholar
Boulanger, J. & Stenhouse, G. B. (2014). The impact of roads on the demography of grizzly bears in Alberta. PLoS ONE 9: e115535.CrossRefGoogle ScholarPubMed
Bunnell, F. E. & Tait, D. E. N. (1980). Bears in models and reality – implications to management. International Conference on Bear Research and Management 3: 15–23.Google Scholar
Bunnell, F. E. & Tait, D. E. N. (1981). Populations dynamics of bears–implications. In: Fowler, C. W. & Smith, T. D. (Eds.), Dynamics of large mammal populations (pp. 75–98). New York, NY: John Wiley & Sons.Google Scholar
Byers, C. R. & Bettas, G. A. (1999). Records of North American big game, 11th edition. Missoula, MT: Boone and Crockett Club.Google Scholar
Campbell, B. H. (1999). Homing of translocated brown bears (Ursus arctos) in coastal south-central Alaska. Northwest Naturalist 80: 22–25.CrossRefGoogle Scholar
Case, R. L. & Buckland, L. (1998). Reproductive characteristics of grizzly bears in the Kugluktuk area, Northwest Territories, Canada. Ursus 10: 41–47.Google Scholar
Cherry, S., Haroldson, M. A., Robison-Cox, J. & Schwartz, C. C. (2002). Estimating total human-caused mortality from reported mortality using data from radio-instrumented grizzly bears. Ursus 13: 175–184.Google Scholar
Chester, C. C., Hilty, J. A. & Hamilton, L. S. (2013). Mountain gloom and mountain glory revisited: a survey of conservation, connectivity, and climate change in mountain regions. Journal of Mountain Ecology 9: 1–34.Google Scholar
Chomel, B. B., Kasten, R. W., Chappuis, G., Soulier, M. & Kikuchi, Y. (1998). Serological survey of selected canine viral pathogens and zooneses in grizzly bears (Ursus arctos horribilis) and bears (Ursus americanus) from Alaska. Revue scientifique et technique 17: 756–766.CrossRefGoogle Scholar
Ciarniello, L. M., Boyce, M. S., Heard, D. C. & Seip, D. R. (2005). Denning behavior and den site selection of grizzly bears along the Parsnip River, British Columbia, Canada. Ursus 16: 47–58.CrossRefGoogle Scholar
Clapham, M. & Kitchin, J. (2016). Social play in wild brown bears of varying age-sex class. Acta Ethologica 19: 181–188.CrossRefGoogle Scholar
Coleman, T. H., Schwartz, C. C., Gunther, K. A. & Creel, S. (2013). Grizzly bear and human interaction in Yellowstone National Park: an evaluation of bear management areas. Journal of Wildlife Management 77: 1311–1320.CrossRefGoogle Scholar
Collins, G. H., Kovach, S. D. & Hinkes, M. T. (2005). Home range and movements of female brown bears in southwestern Alaska. Ursus 16: 181–189.CrossRefGoogle Scholar
Committee on the Status of Endangered Wildlife in Canada (COSEWIC). (2012). Assessment and status report on the grizzly bear, Ursus arctos, in Canada. Ottawa: Committee on the Status of Endangered Wildlife in Canada.Google Scholar
Coogan, S. C. P., Raubenheimer, D., Stenhouse, G. B. & Nielsen, S.E. (2014). Macronutrient optimization and seasonal diet mixing in a large omnivore, the grizzly bear: a geometric analysis. PLoS ONE 9: e105719.CrossRefGoogle Scholar
Costello, C. M., Cain, S. L., Pils, S. R., et al. (2016a). Diet and macronutrient optimization in wild ursids: a comparison of grizzly bears with sympatric and allopatric black bears. PLoS ONE: 11: e0153702.CrossRefGoogle ScholarPubMed
Costello, C. M., Mace, R. D. & Roberts, L.. (2016b). Grizzly bear demographics in the Northern Continental Divide ecosystem, Montana: Research results (2004–2014) and suggested techniques for management of mortality. Helena, MT: Montana Department of Fish, Wildlife and Parks.Google Scholar
Cottrell, W. O., Keel, K. M., Brooks, J. W., Mead, D. G. & Phillips, J. E. (2013). First report of clinical disease associated with canine distemper virus infection in a wild black bear (Ursus americana). Journal of Wildlife Diseases 49: 1024–1027.CrossRefGoogle Scholar
Craighead, F. C. Jr. & Craighead, J. J. (1972). Grizzly prehibernation and denning activities as determined by radiotracking. Wildlife Monographs 32: 1–35.Google Scholar
Craighead, J. J. & Mitchell, J. A. (1982). Grizzly bear. In: Chapman, J. A. & Feldhamer, G. A. (Eds.), Wild mammals of North America (pp. 515–555). Baltimore, MD: Johns Hopkins University Press.Google Scholar
Craighead, J. J., Craighead, F. C. Jr. & Sumner, J. (1976). Reproductive cycles and rates in the grizzly bear, Ursus arctos horribilis, of the Yellowstone Ecosystem. International Conference on Bear Research and Management 3: 337–356.Google Scholar
Craighead, J. J., Sumner, J. S. & Mitchell, J. A. (1995). The grizzly bears of Yellowstone: Their ecology in the Yellowstone ecosystem, 1959–1992. Washington DC: Island Press.Google Scholar
Craighead, L., Paetkau, D., Reynolds, H. V., Vyse, E. R. & Strobeck, C. (1995). Microsatellite analysis of paternity and reproduction in Arctic grizzly bears. Journal of Heredity 86: 255–261.CrossRefGoogle ScholarPubMed
Cristescu, B., Stenhouse, G. B. & Boyce, M. S. (2014). Grizzly bear ungulate consumption and the relevance of prey size to caching and meat sharing. Animal Behaviour 92: 133–142.Google Scholar
Cristescu, B., Stenhouse, G. B. & Boyce, M. S. (2015). Grizzly bear diet shifting in reclaimed mines. Global Ecology and Conservation 4: 207–220.CrossRefGoogle Scholar
Cristescu, B., Stenhouse, G. B. & Boyce, M. S. (2016a). Large omnivore movements in response to surface mining and mine reclamation. Scientific Reports 6: 19177.CrossRefGoogle ScholarPubMed
Cristescu, B., Stenhouse, G. B., Goski, B. & Boyce, M. S. (2016b). Grizzly bear space use, survival, and persistence in relation to human habitation and access. Human–Wildlife Interactions 10: 240–257.Google Scholar
Cross, P. C., van Manen, F. T., Viana, M., et al. (2018). Estimating distemper virus dynamics among wolves and grizzly bears using serology and Bayesian state-space models. Ecology and Evolution 8: 8726–8735.CrossRefGoogle ScholarPubMed
Darling, L. M. (1987). Habitat use by grizzly bear family groups in interior Alaska. International Conference on Bear Research and Management 7: 169–178.Google Scholar
Davison, J., Ho, S. Y. W., Bray, S. C., et al. (2011). Late-Quaternary biogeographic scenarios for the brown bear (Ursus arctos), a wild mammal model species. Quaternary Science Reviews 30: 418–430.CrossRefGoogle Scholar
Deacy, W., Leacock, W., Armstrong, J. B. & Stanford, J. A. (2016). Kodiak brown bears surf the salmon red wave: direct evidence from GPS collared individuals. Ecology 97: 1091–1098.CrossRefGoogle ScholarPubMed
Deacy, W. W., Armstrong, J. B., Leacock, W. B., et al. (2017). Phenological synchronization disrupts trophic interactions between Kodiak brown bear and salmon. Proceedings of the National Academy of Sciences 114: 10432–10437.CrossRefGoogle ScholarPubMed
Deacy, W. W., Leacock, W. Stanford, J. A. & Armstrong, J. B. (2019). Variation in spawning phenology within salmon populations influences landscape-level patterns of brown bear activity. Ecosphere 10: e02575.CrossRefGoogle Scholar
Dittrich, L. & Kronberger, H. (1963). Biologisch-anatomische Untersuchungen über die Fortpflanzungsbiologie der Braunbären (Ursus arctos L.) und anderer Ursiden in Gefangenschaft. Zeitschrift für SäugetierKunde 28: 129–155.Google Scholar
Eberhardt, L. L. (1995). Population trend estimates from reproductive and survival data. In: Knight, L. L. & Blanchard, B. (Eds.), Yellowstone grizzly bear investigations (pp. 13–19). Bozeman, MT: Department of Interior, National Biological Service, Interagency Grizzly Bear Study Team.Google Scholar
Eberhardt, L. L. (2002). A paradigm for population analysis of long-lived vertebrates. Ecology 83: 2841–2854.CrossRefGoogle Scholar
Ebinger, M. R., Haroldson, M. A., van Manen, F. T., et al. (2015). Detecting grizzly bear use of ungulate carcasses using global positioning system telemetry and activity data. Oecologia 181: 695–708.CrossRefGoogle Scholar
Edwards, M. A., Nagy, J. A. & Derocher, A. E. (2009). Low site fidelity and home range drift in a wide-ranging, large Arctic omnivore. Animal Behaviour 77: 23–28.CrossRefGoogle Scholar
Edwards, M. A., Derocher, A. E., Hobson, K. A., Branigan, M. & Nagy, J. A. (2011). Fast carnivores and slow herbivores: differential foraging strategies among grizzly bears in the Canadian Artic. Oecologia 165: 877–889.CrossRefGoogle Scholar
Edwards, M. A., Derocher, A. E. & Nagy, J. A. (2013). Home range size variation in female arctic grizzly bears relative to reproductive status and resource availability. PLoS ONE 8: e68130.Google ScholarPubMed
Egbert, A. L. & Stokes, A. W. (1976). The social behaviour of brown bears on an Alaskan salmon stream. International Conference on Bear Research and Management 3: 41–56.Google Scholar
Erickson, A. W., Moosman, H. W., Hensel, R. J. & Troyer, W. A. (1968). The breeding biology of the male brown bear (Ursus arctos). Zoologica 53: 85–105.Google Scholar
Erlenbach, J. A., Rode, K. D., Raubenheimer, D. & Robbins, C. T. (2014). Macronutrient optimization and energy maximization determine diets of brown bears. Journal of Mammalogy 95: 160–168.CrossRefGoogle Scholar
Ewer, R. F. (1973). The carnivores. Ithaca, NY: Cornell University Press.Google Scholar
Fagen, R. & Fagen, J. (1990). Play behavior of brown bears (Ursus arctos) and human presence at Pack Creek, Admiralty Island, Alaska. International Conference on Bear Research and Management 8: 315–319.Google Scholar
Fagen, R. & Fagen, J. (2004). Juvenile survival and benefits of play behaviour in brown bears, Ursus arctos. Evolutionary Ecology Research 6: 89–102.Google Scholar
Fagen, R. & Fagen, J. (2009). Play behaviour and multi-year juvenile survival in free-ranging brown bears, Ursus arctos. Evolutionary Ecology Research 11: 1–15.Google Scholar
Felicetti, L. A., Robbins, C. T. & Shipley, L. A. (2003). Dietary protein content alters energy expenditure and composition of the mass gain in grizzly bears (Ursus arctos horribilis). Physiological and Biochemical Zoology 76: 256–261.CrossRefGoogle ScholarPubMed
Felicetti, L. A., Schwartz, C. C., Rye, R. O., et al. (2004). Use of naturally occurring mercury to determine the importance of cutthroat trout to Yellowstone grizzly bears. Canadian Journal of Zoology 82: 493–501.CrossRefGoogle Scholar
Ferguson, S. H. & McLoughlin, P. D. (2000). Effect of energy availability, seasonality, and geographic range on brown bear life history. Ecography 23: 193–200.CrossRefGoogle Scholar
Folk, G. E. Jr., Folk, M. A. & Minor, J. J. (1972). Physiological condition of three species of bears in winter dens. International Conference on Bear Research and Management 2:107–124.Google Scholar
Ford, A. T., Rettie, K. & Clevenger, A. P. (2009). Fostering ecosystem function through an international public-private partnership: a case study of wildlife mitigation measures along the Trans-Canada Highway in Banff National Park, Alberta, Canada. International Journal of Biodiversity Science and Management 5: 181–189.CrossRefGoogle Scholar
Fortin, J. K., Schwartz, C. C., Gunther, K. A., et al. (2013a). Dietary adjustability of grizzly bears and American black bears in Yellowstone National Park. Journal of Wildlife Management 77: 270–281.CrossRefGoogle Scholar
Fortin, J. K., Ware, J. V., Jansen, H. T., Schwartz, C. C. & Robbins, C. T. (2013b). Temporal niche switching by grizzly bears but not American black bears in Yellowstone National Park. Journal of Mammalogy 94: 833–844.CrossRefGoogle Scholar
Frechette, J.-L. & Rau, M. E. (1978). Seasonal changes in the prevalence of ova of Diphyllobothrium ursi and Baylisascaris transfuga in the feces of the black bear (Ursus americanus). Journal of Wildlife Diseases 14: 342–344.CrossRefGoogle ScholarPubMed
French, S. P., French, M. G. & Knight, R. R. (1994). Grizzly bear use of army cutworm moths in the Yellowstone Ecosystem. International Conference on Bear Research and Management 9: 389–399.Google Scholar
Gardner, C. L., Pamperin, N. J. & Benson, J. F. (2014). Movement patterns and space use of maternal grizzly bears influence cub survival in Interior Alaska. Ursus 25: 121–138.CrossRefGoogle Scholar
Garshelis, D. L., Gibeau, M. L. & Herrero, S. (2005). Grizzly bear demographics in and around Banff National Park and Kananaskis Country, Alberta. Journal of Wildlife Management 69: 277–297.2.0.CO;2>CrossRefGoogle Scholar
Gau, R. J. (1998). Food habitats, body condition, and habitat of the barren-ground grizzly bear. MS thesis, University of Alaska.Google Scholar
Gau, R. J., Kutz, S. & Elkin, B. T. (1999). Parasites in grizzly bears from the central Canadian Arctic. Journal of Wildlife Diseases 35: 618–621.CrossRefGoogle ScholarPubMed
Gau, R. J., Case, R., Penner, D. F. & McLoughlin, P. D. (2002). Feeding patterns of barren-ground grizzly bears in the central Canadian Arctic. Arctic 55: 339–344.CrossRefGoogle Scholar
Gau, R. J., McLoughlin, P. D., Case, R., et al. (2004). Movements of subadult male grizzly bears, Ursus arctos, in the Central Canadian Arctic. Canadian Field Naturalist 118: 239–242.CrossRefGoogle Scholar
Gende, S. M. & Quinn, T. P. (2004). The relative importance of prey density and social dominance in determining energy intake by bears feeding on Pacific salmon. Canadian Journal of Zoology 82: 75–85.CrossRefGoogle Scholar
Gibeau, M. L., Clevenger, A. P., Herrero, S. & Wierzchowski, J. (2002). Grizzly bear response to human developments and activities in the Bow River Watershed, Alberta, Canada. Biological Conservation 103: 227–236.CrossRefGoogle Scholar
Glass, B. P. (1974). A key to the skulls of North American mammals, 2nd edition. Stillwater, OK: Author.Google Scholar
Glenn, L. P. & Miller, L. H. (1980). Seasonal movements of an Alaska Peninsula brown bear population. International Conference for Bear Research and Management 4: 307–312.Google Scholar
Goldstein, M. I., Poe, A. J., Suring, L. H., Nielson, R. M. & McDonald, T. L. (2010). Brown bear den habitat and winter recreation in south-central Alaska. Journal of Wildlife Management 74: 35–42.CrossRefGoogle Scholar
Gordon, K. R. (1977). Molar measurements as a taxonomic tool in Ursus. Journal of Mammalogy 58: 247–248.CrossRefGoogle Scholar
Graham, K. & Stenhouse, G. B. (2014). Home range, movements, and denning chronology of the grizzly bear (Ursus arctos) in West-central Alberta. Canadian Field-Naturalist 128: 223–234.CrossRefGoogle Scholar
Graves, T. A., Farley, S. & Servheen, C. (2006). Frequency and distribution of highway crossings by Kenai Peninsula brown bears. Wildlife Society Bulletin 34: 800–808.CrossRefGoogle Scholar
Guilday, J. E. (1968). Grizzly bears from eastern North America. American Midland Naturalist 79: 247–50.CrossRefGoogle Scholar
Gunther, K. A. & Smith, D. W. (2004). Interactions between wolves and female grizzly bears with cubs in Yellowstone National Park. Ursus 15: 232–238.2.0.CO;2>CrossRefGoogle Scholar
Gunther, K. A., Shoemaker, R. R., Frey, K. L., et al. (2014). Dietary breadth of grizzly bears in the Greater Yellowstone Ecosystem. Ursus 25: 60–72.CrossRefGoogle Scholar
Hall, E. R. (1984). Geographic variation among brown and grizzly bears (Ursus arctos) in North America (Special Publication 13). Lawrence, KS: Museum of Natural History, University of Kansas.Google Scholar
Hamer, D. & Herrero, S. (1987). Wildfire’s influence on grizzly bear feeding ecology in Banff National Park, Alberta. International Conference on Bear Research and Management 7: 179–186.Google Scholar
Hamer, D. & Herrero, S. (1990). Courtship and use of mating areas by grizzly bears in the Front Ranges of Banff National Park, Alberta. Canadian Journal of Zoology 68: 2695–2697.CrossRefGoogle Scholar
Harding, L. E. (1976). Den-site characteristics of arctic coastal grizzly bears (Ursus zrctos) on Richards Island, Northwest Territories, Canada. Canadian Journal of Zoology 54: 1357–1363.CrossRefGoogle Scholar
Harestad, A. S. & Bunnell, F. L. (1979). Home range and body weight – a reevaluation. Ecology 60: 389–402.CrossRefGoogle Scholar
Haroldson, M. A., Ternent, M. A., Gunther, K. A. & Schwartz, C. C. (2002). Grizzly bear denning chronology and movements in the Greater Yellowstone Ecosystem. Ursus 13: 29–37.Google Scholar
Haroldson, M. A., Schwartz, C. C., Kendall, K. C., et al. (2010). Genetic analysis of individual origins supports isolation of grizzly bears in the Greater Yellowstone Ecosystem. Ursus 21: 1–13.CrossRefGoogle Scholar
Haroldson, M. A., Gunther, K. A., Cain, S. L., Wilmot, K. R. & Wyman, T. (2015). Grizzly cub adoptions confirmed in Yellowstone and Grand Teton National Parks. Yellowstone Science 23: 58–61.Google Scholar
Hechtel, J. L. (1985). Activity and food habits of barren-ground grizzly bears in Arctic Alaska. MS thesis, University of Montana.Google Scholar
Herrero, S. (1972). Aspects of evolution and adaptation in American black bears (Ursus americanus Pallas) and brown and grizzly bears (U. arctos Linné.) of North America. International Conference for Bear Research and Management 2: 221–231.Google Scholar
Herrero, S., Smith, T., Debruyn, T. D., Gunther, K. A. & Matt, C. A. (2005). From the field: brown bear habituation to people – safety, risks, and benefits. Wildlife Society Bulletin 33: 362–373.CrossRefGoogle Scholar
Hessing, P. & Aumiller, L. (1994). Observations of conspecific predation by brown bears, Ursus arctos, in Alaska. Canadian Field-Naturalist 108: 332–336.Google Scholar
Hilderbrand, G. V., Jenkins, S. G., Schwartz, C. C., Hanley, T. A. & Robbins, C. T. (1999a). Effect of seasonal differences in dietary meat intake on changes in body mass and composition in wild and captive brown bears. Canadian Journal of Zoology 77: 1623–1630.CrossRefGoogle Scholar
Hilderbrand, G. V., Schwartz, C. C., Robbins, C. T., et al. (1999b). The importance of meat, particularly salmon, to body size, population productivity, and conservation of North America brown bears. Canadian Journal of Zoology 77: 132–138.CrossRefGoogle Scholar
Hilderbrand, G. V., Schwartz, C. C., Robbins, C. T. & Hanley, T. A. (2000). Effect of hibernation and reproductive status on body mass and condition of coastal brown bears. Journal of Wildlife Management 64: 178–183.CrossRefGoogle Scholar
Hovey, F. W. & McLellan, B. N. (1996). Estimating population growth of grizzly bears from the Flathead River drainage using computer simulations of reproduction and survival rates. Canadian Journal of Zoology 74: 1409–1416.CrossRefGoogle Scholar
Interagency Grizzly Bear Study Team. (2012). Updating and evaluating approaches to estimate population size and sustainable mortality limits for grizzly bears in the Greater Yellowstone Ecosystem. Bozeman, MT: US Geological Survey, Northern Rocky Mountain Science Center, Interagency Grizzly Bear Study Team.Google Scholar
Jimenez, M. D., Asher, V. J., Bergman, C., Bangs, E. E. & Woodruff, S. P. (2008). Gray wolves, Canis lupus, killed by cougars, Puma concolor, and a grizzly bear, Ursus arctos, in Montana, Alberta, and Wyoming. Canadian Field Naturlist 122: 76–78.CrossRefGoogle Scholar
Judd, S. L., Knight, R. R. & Blanchard, B. M. (1986). Denning of grizzly bears in the Yellowstone National Park area. International Conference on Bear Research and Management 6: 111–117.Google Scholar
Kasworm, W. F., Proctor, M. F., Servheen, C. & Paetkau, D. (2007). Success of grizzly bear population augmentation in northwest Montana. Journal of Wildlife Management 71: 1261–1266.CrossRefGoogle Scholar
Kasworm, W. F., Radandt, T. G., Teisberg, J. E., et al. (2018). Cabinet-Yaak grizzly bear recovery area 2017 research and monitoring progress report. Missoula, MT: US Fish and Wildlife Service.Google Scholar
Keay, J. A., Robbins, C. T. & Farley, S. D. (2018). Characteristics of a naturally regulated grizzly bear population. Journal of Wildlife Management 82: 789–801.CrossRefGoogle Scholar
Kendall, K. C., Stetz, J. B., Roon, D. A., et al. (2008). Grizzly bear density in Glacier National Park, Montana. Journal of Wildlife Management 72: 1693–1705.CrossRefGoogle Scholar
Kendall, K. C., Macleod, A. C., Boyd, K. L., et al. (2015). Density, distribution, and genetic structure of grizzly bears in the Cabinet-Yaak Ecosystem. Journal of Wildlife Management 80: 314–331.CrossRefGoogle Scholar
Kingsley, M. C. S., Nagy, J. A. & Russell, R. H. (1983). Patterns of weight gain and loss for grizzly bears in northern Canada. International Conference for Bear Research and Management 5: 174–178.Google Scholar
Kite, R., Nelson, T., Stenhouse, G. B. & Darimont, C. (2016). A movement-driven approach to quantifying grizzly bear (Ursus arctos) near-road movement patterns in west-central Alberta, Canada. Biological Conservation 195: 24–32.CrossRefGoogle Scholar
Klinka, D. R. & Reimchen, T. E. (2002). Nocturnal and diurnal foraging behavior of brown bears (Ursus arctos) on a salmon stream in coast British Columbia. Canadian Journal of Zoology 80: 1317–1322.CrossRefGoogle Scholar
Knight, R. R., Blanchard, B. & Kendall, K. (1981). Yellowstone grizzly bear investigations: Annual report of the Interagency Study Team, 1980. Bozeman, MT: US Geological Survey, Northern Rocky Mountain Science Center, Interagency Grizzly Bear Study Team.Google Scholar
Knowles, S., Bodenstein, B. L., Hamon, T., Saxton, M. W. & Hall, J. S. (2018). Infectious canine hepatitis in a brown bear (Ursus arctos horribilis) from Alaska. Journal of Wildlife Diseases 54: 642–645.CrossRefGoogle Scholar
Koch, E. (1941). Big game in Montana from early historical records. Journal of Wildlife Management 5: 357–370.CrossRefGoogle Scholar
Koene, P., Ardesch, J., Ludriks, A., et al. (2002). Interspecific and intraspecific social interactions among brown bears and wolves in an enclosure. Ursus 13: 85–93.Google Scholar
Kovach, S. D., Collins, G. H., Hinkes, M. T. & Denton, J. W. (2006). Reproduction and survival of brown bears in southwest Alaska, USA. Ursus 17: 16–29.CrossRefGoogle Scholar
Kumar, V., Lammers, F., Bidon, T., et al. (2017). The evolutionary history of bears is characterized by gene flow across species. Scientific Reports 7: 46487.CrossRefGoogle ScholarPubMed
Kurtén, B. (1973). Transberingian relationships of Ursus arctos Linne (brown and grizzly bears). Commentation Biology 65: 1–10.Google Scholar
Ladle, A., Avgar, T., Wheatley, M., et al. (2019). Grizzly bear response to spatio‐temporal variability in human recreational activity. Journal of Applied Ecology 56: 375–386.CrossRefGoogle Scholar
Lamb, C. T., Mowat, G., Reid, A., et al. (2018). Effects of habitat quality and access management on the density of a recovering grizzly bear population. Journal of Applied Ecology 55: 1406–1417.CrossRefGoogle Scholar
LeFranc, M. N. Jr., Moss, M. B., Patnode, K. A. & Sugg, W. C. (1987). Grizzly bear compendium. Washington, DC: Interagency Grizzly Bear Committee.Google Scholar
Lewis, T. M. & Lafferty, D. J. R. (2014). Brown bears and wolves scavenge humpback whale carcass in Alaska. Ursus 25: 8–13.CrossRefGoogle Scholar
Linnell, J. D. C., Swenson, J. E., Anderson, R. & Barnes, B. (2000). How vulnerable are denning bears to disturbance? Wildlife Society Bulletin 28: 400–413.Google Scholar
López-Alfaro, C., Coogan, S. C. P., Robbins, C. T., Fortin, J. K. & Nielsen, S. E. (2015). Assessing nutritional parameters of brown bear diets among ecosystems gives insight into differences among populations. PLoS ONE 10: e0128088.CrossRefGoogle ScholarPubMed
Mace, R. D. & Waller, J. S. (1997). Grizzly bear ecology in the Swan Mountains. Final Report. Helena, MT: Montana Department of Fish, Wildlife and Parks.Google Scholar
Mace, R. D. & Waller, J. S. (1998). Demography and population trend of grizzly bears in the Swan Mountains, Montana. Conservation Biology 12: 1005–1016.CrossRefGoogle Scholar
Mace, R. D., Waller, J. S., Manley, T. L., Lyon, L. J. & Zuring, H. (1996). Relationships among grizzly bears, roads, and habitat use in the Swan Mountains, Montana. Journal of Applied Ecology 33: 1395–1404.CrossRefGoogle Scholar
Mace, R. D., Carney, D. W., Chilton-Radandt, T., et al. (2012). Grizzly bear population vital rates and trend in the Northern Continental Divide Ecosystem, Montana. Journal of Wildlife Management 76: 119–128.CrossRefGoogle Scholar
MacHutchon, A. G. & Wellwood, D. W. (2003). Grizzly bear food habits in the northern Yukon, Canada. Ursus 14: 225–235.Google Scholar
MacHutchon, A. G., Himmer, S. & Bryden, C. A. (1993). Khutzeymateen Valley grizzly bear study. Wildlife Habitat Research Report No. 31. Victoria: British Columbia Ministry of Forests.Google Scholar
MacHutchon, A. G., Himmer, S., Davis, H. & Gallagher, M. (1998). Temporal and spatial activity patterns among coastal bear populations. Ursus 10: 539–546.Google Scholar
MacPherson, A. H. (1965). The barren-ground grizzly bear and its survival in Canada. Canadian Audubon 27: 2–8.Google Scholar
Mangipane, L. S., Belant, J. L., Hiller, T. L., et al. (2018) Influences of landscape heterogeneity on home-range sizes of brown bears. Mammalian Biology 88: 1–7.CrossRefGoogle Scholar
García Marín, J. F., Royo, L. J., Oleaga, A., et al. (2018). Canine adenovirus type 1 (CAdV‐1) in free‐ranging European brown bear (Ursus arctos arctos): a threat for Cantabrian population? Transboundary and Emerging Diseases 65: 2049–2056.CrossRefGoogle ScholarPubMed
Matson, G. M., Van Daele, L., Goodwin, E., et al. (1993). A laboratory manual for cementum age determination of Alaskan brown bear first premolar teeth. Milltown, MT: Matson’s Laboratory.Google Scholar
Mattson, D. (1997). Use of ungulates by Yellowstone grizzly bears Ursus arctos. Biological Conservation 81: 161–177.CrossRefGoogle Scholar
Mattson, D. (2004). Exploitation of pocket gophers and their food caches by grizzly bears. Journal of Mammalogy 85: 731–742.CrossRefGoogle Scholar
Mattson, D. J. & Merrill, T. (2002). Extirpations of grizzly bears in the contiguous United States, 1850–2000. Conservation Biology 16: 1123–1136.CrossRefGoogle Scholar
Mattson, D. J., Knight, R. R. & Blanchard, B. M. (1992). Cannibalism and predation on black bears by grizzly bears in the Yellowstone Ecosystem, 1975–1990. Journal of Mammalogy 73: 422–425.CrossRefGoogle Scholar
Mattson, D. J., Wright, R. G., Kendall, K. C. & Martinka, C. J. (1995). Grizzly bears. In: LaRoe, E. T., Farris, G. S., Puckett, C. E., Doran, P. D. & Mac, M. J. (Eds.), Our living resources: A report to the nation on the distribution, abundance, and health of US plants, animals, and ecosystems (pp. 103–105). Washington, DC: US Department of the Interior, National Biological Service.Google Scholar
McLellan, B. N. (1989). Dynamics of a grizzly bear population during a period of industrial resource extraction I. Density and age-sex composition. Canadian Journal of Zoology 67: 1857–1868.Google Scholar
McLellan, B. N. (1994). Density-dependent population regulation of brown bears. In Taylor, M. (Ed.), Density-dependent population regulation in black, brown, and polar bears (pp. 15–24). International Association for Bear Research and Management Monograph Series No. 3.Google Scholar
McLellan, B. N. (1998). Maintaining viability of brown bears along the southern fringe of their distribution. Ursus 10: 607–611.Google Scholar
McLellan, B. N. (2005). Sexually selected infanticide in grizzly bears: the effects of hunting on cub survival. Ursus 16: 141–156.CrossRefGoogle Scholar
McLellan, B. N. (2011). Implications of high-energy and low-protein diet on the body composition, fitness, and competitive ability of black (Ursus americanus) and grizzly (Ursus arctos) bears. Canadian Journal of Zoology 89: 546–558.CrossRefGoogle Scholar
McLellan, B. N. (2015). Some mechanisms underlying variation in vital rates of grizzly bear on a multiple-use landscape. Journal of Wildlife Management 79: 749–765.CrossRefGoogle Scholar
McLellan, B. N. & Banci, V. (1999). Status and management of the brown bear in Canada. In: Servheen, C., Herrero, S. & Peyton, B. (Eds.), Bears: Status survey and conservation action plan (pp. 46–50). Gland: IUCN/SSC Bear and Polar Bear Specialist Groups, International Union for the Conservation of Nature.Google Scholar
McLellan, B. N. & Hovey, F. W. (1995). The diet of grizzly bears in the Flathead River drainage of southeast British Columbia. Canadian Zoology of Journal 73: 704–712.CrossRefGoogle Scholar
McLellan, B. N. & Hovey, F. W. (2001). Habitats selected by grizzly bears in a multiple use landscape. Journal of Wildlife Management 65: 92–99.Google Scholar
McLellan, B. N. & Reiner, D. C. (1994). A review of bear evolution. Ursus 9: 85–96.Google Scholar
McLellan, B. N., Hovey, F. W., Mace, R. D., et al. (1999). Rates and causes of grizzly bear mortality in the interior mountains of British Columbia, Alberta, Montana, Washington, and Idaho. Journal of Wildlife Management 63: 911–920.CrossRefGoogle Scholar
McLellan, B. N., Mowat, G., Hamilton, T. & Hatter, I. (2017a). Sustainability of the grizzly bear hunt in British Columbia, Canada. Journal of Wildlife Management 81: 218–229.CrossRefGoogle Scholar
McLellan, B. N., Proctor, M. F., Huber, Đ. & Michel, S. (2017b). Ursus arctos. IUCN Red List of Threatened Species, www.iucnredlist.org/details/41688/0 (accessed April 10, 2019).Google Scholar
McLellan, M. L. & McLellan, B. N. (2015). Effect of season and high ambient temperature on activity levels and patterns of grizzly bears (Ursus arctos). PLoS ONE 10: e0117734.CrossRefGoogle ScholarPubMed
McLoughlin, P. D. (2000). The spatial organization and habitat selection patterns of barren-ground grizzly bears in the central Arctic. PhD dissertation, University of Saskatchewan.Google Scholar
McLoughlin, P. D., Case, R. L., Gau, R. J., Ferguson, S. H. & Messier, F. (1999). Annual and seasonal movement patterns of barren-ground grizzly bears in the central Northwest Territories. Ursus 11: 79–86.Google Scholar
McLoughlin, P. D., Ferguson, S. H. & Messier, F. (2000). Intraspecific variation in home range overlap with habitat quality: a comparison among brown bear populations. Evolutionary Ecology 14: 39–60.CrossRefGoogle Scholar
McLoughlin, P. D., Case, R. L., Gau, R. J., et al. (2002a). Hierarchical habitat selection by barren-ground grizzly bears in the central Canadian arctic. Oecologia 132: 102–108.CrossRefGoogle ScholarPubMed
McLoughlin, P. D., Cluff, H. D. & Messier, F. (2002b). Denning ecology of barren-ground grizzly bears in the central Arctic. Journal of Mammalogy 83: 188–198.2.0.CO;2>CrossRefGoogle Scholar
McLoughlin, P. D., Cluff, H. D., Gau, R. J., et al. (2003a). Effect of spatial differences in habitat on home ranges of grizzly bears. Ecoscience 10: 11–16.CrossRefGoogle Scholar
McLoughlin, P. D., Taylor, M. K., Cluff, H. D., et al. (2003b). Demography of barren-ground grizzly bears. Canadian Journal of Zoology 81: 294–301.CrossRefGoogle Scholar
McNab, B. K. (1963). Bioenergetics and the determination of home range size. American Naturalist 97: 133–140.CrossRefGoogle Scholar
Merriam, C. H. (1918). Review of the grizzly and big brown bears of North America (genus Ursus) with the description of a new genus, Vetularctos. North American Fauna 41: 1–136.CrossRefGoogle Scholar
Milakovic, B. & Parker, K. L. (2012). Quantifying carnivory by grizzly bears in a multi-ungulate system. Journal of Wildlife Management 77: 39–47.CrossRefGoogle Scholar
Miller, C. R., Waits, L. P. & Joyce, P. (2006). Phylogeography and mitochondrial diversity of extirpated brown bear (Ursus arctos) populations in the contiguous United States and Mexico. Molecular Ecology 15: 4477–4485.CrossRefGoogle ScholarPubMed
Miller, S. D. (1990). Denning ecology of brown bears in south-central Alaska and comparisons with sympatric black bear population. International Conference for Bear Research and Management 8: 279–287.Google Scholar
Miller, S. D. (1993). Brown bears in Alaska: A statewide management overview. Wildlife Technical Bulletin 11. Juneau, AK: Alaska Department of Fish and Game.Google Scholar
Miller, S. D. & Ballard, W. B. (1982). Homing of transplanted Alaskan brown bears. Journal of Wildlife Management 46: 869–876.CrossRefGoogle Scholar
Miller, S. D. & Schoen, J. (1999). Status and management of the brown bear in Alaska. In: Servheen, C., Herrero, S. & Peyton, B. (Eds.), Bears: Status survey and conservation action plan (pp. 40–46). Gland: IUCN/SSC Bear and Polar Bear Specialist Groups, International Union for the Conservation of Nature.Google Scholar
Miller, S. D. & Sellers, R. A. (1992). Brown bear density on the Alaska Peninsula at Black Lake, Alaska. Final report on completion of density estimation objectives of cooperative interagency brown bear studies on the Alaska Peninsula. Juneau, AK: Alaska Department of Fish and Game.Google Scholar
Miller, S. D. & Tutterow, V. L. (1999). Characteristics of nonsport mortalities to brown and black bears and human injuries from bears in Alaska. Ursus 11: 239–250.Google Scholar
Miller, S. D., Sellers, R. A. & Keay, J. A. (2003). Effects of hunting on brown bear cub survival and litter size in Alaska. Ursus 14: 130–152.Google Scholar
Miller, S. D., Schoen, J. W., Faro, J. & Klein, D. R. (2011). Trends in intensive management of Alaska’s grizzly bears, 1980–2010. Journal of Wildlife Management 75: 1243–1252.CrossRefGoogle Scholar
Miller, S. D., McLellan, B. N. & Derocher, A. E. (2013). Conservation and management of large carnivores in North America. International Journal of Environmental Studies 70: 383–398.CrossRefGoogle Scholar
Miller, S. D., Wilder, J. & Wilson, R. R. (2015). Polar bear-grizzly bear interactions during the autumn open-water period in Alaska. Journal of Mammalogy 96: 1317–1325.CrossRefGoogle Scholar
Miller, S. D., Schoen, J. W. & Schwartz, C. C. (2017). Tends in brown bear reduction efforts in Alaska, 1980–2017. Ursus 28:135–149.CrossRefGoogle Scholar
Milligan, S., Brown, L., Hobson, D., Frame, P. & Stenhouse, G. (2018). Factors affecting the success of grizzly bear translocations. Journal of Wildlife Management 82: 519–530CrossRefGoogle Scholar
Modafferi, R. D. (1984). Review of Alaska Peninsula brown bear investigations. Final Report Federal Aid in Wildlife Restoration Projects W-17-10, W-17-11, W-21-1, W-21-2, W-22-1. Juneau, AK: Alaska Department of Fish and Game.Google Scholar
Morehouse, A. T., Graves, T. A., Mikle, N. & Boyce, M. S. (2016). Nature vs. nurture: evidence for social learning of conflict behaviour in grizzly bears. PLoS ONE 11: e0165425.CrossRefGoogle ScholarPubMed
Mowat, G. & Heard, D. C. (2006). Major components of grizzly bear diet across North America. Canadian Journal of Zoology 84: 473–489.CrossRefGoogle Scholar
Mowat, G., Heard, D. C. & Schwarz, C. J. (2013). Predicting grizzly bear density in western North America. PLoS ONE 8: e82757.CrossRefGoogle ScholarPubMed
Munro, R. H. M., Nielsen, S. E., Price, M. H., Stenhouse, G. B. & Boyce, M. S. (2006). Seasonal and diel patterns of grizzly bear diet and activity in west-central Alberta. Journal of Mammalogy 87: 1112–1121.CrossRefGoogle Scholar
Nagy, J. A. & Haroldson, M. A. (1989). Comparison of some home range and population parameters among four grizzly bear populations in Canada. International Conference for Bear Research and Management 8: 227–235.Google Scholar
Nagy, J. A. & Russell, R. H. (1978). Ecological studies of the boreal forest grizzly bear Ursus arctos: annual report for 1977. Edmonton: Canadian Wildlife Service.Google Scholar
Nagy, J. A., Russell, R. H., Pearson, A. M., Kingsley, M. C. & Goski, B. C. (1983). Ecological studies of the grizzly bear in arctic mountains, northern Yukon Territories, 1972 to 1975. Edmonton: Canadian Wildlife Service.Google Scholar
Nelson, R. A. (1980). Protein and fat metabolism in hibernating bears. Federation Proceedings 39: 2955–2958.Google ScholarPubMed
Nelson, R. A., Folk, G. E. Jr., Pfeiffer, W. W., et al. (1983). Behavior, biochemistry, and hibernation in black, grizzly, and polar bears. Proceedings Eastern Workshop on Black Bear Research and Management 7: 48–53.Google Scholar
Nevin, O. T. & Gilbert, B. K. (2005). Measuring the cost of risk avoidance in brown bears: further evidence of positive impacts of ecotourism. Biological Conservation 121: 611–622.CrossRefGoogle Scholar
Nielsen, S. E., Boyce, M. S., Stenhouse, G. B. & Munro, R. H. M. (2003). Development and testing of phenologically driven grizzly bear habitat models. Ecoscience 10: 1–10.CrossRefGoogle Scholar
Nielsen, S. E., Herrero, S., Boyce, M. S., et al. (2004). Modeling the spatial distribution of human-caused grizzly bear mortalities in the Central Rockies Ecosystem of Canada. Biological Conservation 120: 101–113.CrossRefGoogle Scholar
Nielsen, S. E., Larsen, T. A., Stenhouse, G. B. & Coogan, S. C .P. (2017). Complementary food resources of carnivory and frugivory affect local abundance of an omnivorous carnivore. Oikos 126: 369–380.CrossRefGoogle Scholar
Northrup, J. M., Pitt, J., Muhly, T. B., et al. (2012a). Vehicle traffic shapes grizzly bear behaviour on a multiple‐use landscape. Journal of Applied Ecology 49: 1159–1167.CrossRefGoogle Scholar
Northrup, J. M., Stenhouse, G. B. & Boyce, M. S. (2012b). Agricultural lands as ecological traps for grizzly bears. Animal Conservation 15: 369–377.CrossRefGoogle Scholar
Oftedal, O. T. & Gittleman, J. L. (1989). Patterns of energy output during reproduction in carnivores. In: Gittleman, J. L. (Ed.), Carnivore behavior, ecology, and evolution (pp. 355–378). Boston, MA: Springer.CrossRefGoogle Scholar
Paetkau, D., Shields, G. F. & Strobeck, C. (1998a). Gene flow between insular, coastal and interior populations of brown bears in Alaska. Molecular Ecology 7: 1283–1292.CrossRefGoogle ScholarPubMed
Paetkau, D., Waits, L. P., Clarkson, P. L., et al. (1998b). Variation in genetic diversity across the range of North American brown bears. Conservation Biology 12: 418–429.CrossRefGoogle Scholar
Palmer, K. (2015). Oldest brown bear in captivity in North America dies at Ohio zoo. Thomson Reuters. www.reuters.com/article/us-usa-ohio-bear/oldest-brown-bear-in-captivity-in-north-america-dies-at-ohio-zoo-idUSKCN0RB1UG20150911 (accessed May 29, 2019).Google Scholar
Pasitschniak-Arts, M. (1993). Ursus arctos. Mammalian Species 439: 1–10.CrossRefGoogle Scholar
Pearson, A. M. (1975). The northern interior grizzly bear Ursus arctos L. Report Series No. 34. Ottawa: Canadian Wildlife Service.Google Scholar
Peck, C. P., van Manen, F. T., Costello, C. M., et al. (2017). Potential paths for male‐mediated gene flow to and from an isolated grizzly bear population. Ecosphere 8: e01969.CrossRefGoogle Scholar
Peterson, R. L. (1965). A well-preserved grizzly bear skull recovered from a late glacial deposit near Lake Simcoe, Ontario. Nature 208: 1233–1234.CrossRefGoogle Scholar
Pigeon, K. E., Nielsen, S. E., Stenhouse, G. B. & Coté, S. D. (2014). Den selection by grizzly bears on a managed landscape. Journal of Mammalogy 95: 559–571.CrossRefGoogle Scholar
Pigeon, K. E., Coté, S. D. & Stenhouse, G. B. (2016a). Assessing den selection and den characteristics of grizzly bears. Journal of Wildlife Management 80: 884–893.CrossRefGoogle Scholar
Pigeon, K. E., Stenhouse, G. B. & Coté, S.D. (2016b). Drivers of hibernation: linking food and weather to denning behaviour of grizzly bears. Behavioral Ecology and Sociobiology 70: 1745–1754.CrossRefGoogle Scholar
Podruzny, S. R., Cherry, S., Schwartz, C. C. & Landenburger, L. A. (2002). Grizzly bear denning and potential conflict areas in the Greater Yellowstone Ecosystem. Ursus 13: 19–28.Google Scholar
Proctor, M. F. & MacHutchon, G. (2019). Grizzly bear management and habitat use in the agricultural Creston Valley of southeast British Columbia and a vision for long-term management. Trans-border Grizzly Bear Project. Kaslo: Trans-border Grizzly Bear Project.Google Scholar
Proctor, M. F., McLellan, B. N., Strobeck, C. & Barclay, R. M. (2004). Gender-specific dispersal distance of grizzly bears estimated by genetic analysis. Canadian Journal of Zoology 82: 1108–1118.CrossRefGoogle Scholar
Proctor, M. F., Paetkau, D., McLellan, B. N., et al. (2012). Population fragmentation and inter-ecosystem movements of grizzly bears in Western Canada and the Northern United States. Wildlife Monographs 180: 1–46.CrossRefGoogle Scholar
Proctor, M. F., Nielsen, S. E., Kasworm, W. F., et al. (2015). Grizzly bear connectivity mapping in the Canada–US trans-border region. Journal of Wildlife Management 79: 544–558.CrossRefGoogle Scholar
Proctor, M. F., Lamb, C. T. & MacHutchon, A. G. (2017). The grizzly dance of berries and bullets: The relationship between bottom up food resources, huckleberries, and top down mortality risk on grizzly bear population processes in southeast British Columbia. Kaslo: Trans-border Grizzly Bear Project.Google Scholar
Proctor, M. F., Kasworm, W. F., Annis, K. M., et al. (2018a). Conservation of threatened Canada-USA trans-border grizzly bears linked to comprehensive conflict reduction. Human–Wildlife Interactions 12: 348–372.Google Scholar
Proctor, M. F., McLellan, B. N., Stenhouse, G. B., et al. (2018b). Resource roads and grizzly bears in British Columbia and Alberta. Canadian Grizzly Bear Management Series, Resource Road Management. Kaslo: Trans-border Grizzly Bear Project.Google Scholar
Quimby, R. & Snarski, D. J. (1974). A study of fur-bearing mammals associated with gas pipeline routes in Alaska. In: Jakimchuk, R. D. (Ed.), Distribution of moose, sheep, muskox and fur-bearing mammals in northeastern Alaska (pp. 11–36). Arctic Gas Biological Report Series, Volume 6. Canadian Arctic Gas Study Ltd. and Alaskan Arctic Gas Study Company.Google Scholar
Quinn, T. P., Gende, S. M., Ruggerone, G. T. & Rogers, D. E. (2003). Density-dependent predation by brown bears (Ursus arctos) on sockeye salmon (Oncorhynchus nerka). Canadian Journal of Fisheries and Aquatic Sciences 60: 553–562.CrossRefGoogle Scholar
Rausch, R. L. (1953). On the status of some arctic mammals. Arctic 6: 91–148.CrossRefGoogle Scholar
Rausch, R. L. (1954). Studies on the helminth fauna of Alaska. XXI. Taxonomy, morphological variation, and ecology of Diphyllobothrium ursi n. sp. provis. on Kodiak Island. Journal of Parasitology 40: 540–563.CrossRefGoogle ScholarPubMed
Rausch, R. L. (1963). Geographic variation in size of North American brown bears, Ursus arctos L., as indicated by condylobasal length. Canadian Journal of Zoology 41: 33–45.CrossRefGoogle Scholar
Rausch, R. L. (1969). Morphogenesis and age-related structure of permanent canine teeth in the brown bear, Ursus arctos L., in Arctic Alaska. Zeitschrift für Morphologie der Tiere 66: 167–188.CrossRefGoogle Scholar
Reynolds, H. V. (1976). North Slope grizzly bear studies. Federal Aid in Wildlife Restoration Final Report for Projects W-17-6 and W-17-7, Jobs 4.8R, 4.9R, 4.10R, and 4.11R. Juneau, AK: Alaska Department of Fish and Game.Google Scholar
Reynolds, H. V., Curatolo, J.A. & Quimby, R. (1976). Denning ecology of grizzly bears in Northeastern Alaska. International Conference on Bear Research and Management 3: 403–409.Google Scholar
Reynolds, P. E., Reynolds, H. V. & Shideler, R. T. (2002). Predation and multiple kills of muskoxen by grizzly bears. Ursus 13: 79–84.Google Scholar
Riley, S. J., Aune, K., Mace, R. D. & Madel, M. J. (2004). Translocation of nuisance grizzly bears in northwestern Montana. International Conference on Bear Research and Management 9: 567–573.Google Scholar
Ripple, W. J., Miller, S. D., Schoen, J. W. & Rabinowitch, S. P. (2019). Large carnivores under assault in Alaska. PLoS Biology 17: e3000090.CrossRefGoogle ScholarPubMed
Robbins, C. T., Schwartz, C. C. & Felicetti, L. A. (2004). Nutritional ecology of ursids: a review of newer methods and management implications. Ursus 15: 161–171.2.0.CO;2>CrossRefGoogle Scholar
Robbins, C. T., Fortin, J. K., Rode, K. D., Shipley, L. A. & Felicetti, L. A. (2007). Optimizing protein intake as a foraging strategy to maximize mass gain in an omnivore. Oikos 116: 1675–1682.CrossRefGoogle Scholar
Robbins, C. T., Ben-David, M., Fortin, J. K. & Nelson, O. L. (2012). Maternal condition determines birth date and growth of newborn bear cubs. Journal of Mammalogy 93: 540–546.CrossRefGoogle Scholar
Robbins, C. T., Woodford, N. L., Clyde, G. G., et al. (2018). Salmon poisoning disease in grizzly bears with population recovery implications. Journal of Wildlife Management 82:1396–1402.CrossRefGoogle Scholar
Rockwell, D. (1991). Giving voice to bear. Niwot, CO: Roberts Rinehart publishers.Google Scholar
Rode, K. D. & Robbins, C. T. (2000). Why bears consume mixed diets during fruit abundance. Canadian Journal of Zoology 78: 1640–1645.CrossRefGoogle Scholar
Rode, K. D., Robbins, C. T. & Shipley, L. A. (2001). Constraints on herbivory by grizzly bears. Oecologia 128: 62–71.CrossRefGoogle ScholarPubMed
Rode, K. D., Farley, S. D., Fortin, J. & Robbins, C. T. (2006a). Nutritional consequences of experimentally introduced tourism in brown bears. Journal of Wildlife Management 71: 929–939.CrossRefGoogle Scholar
Rode, K. D., Farley, S. D. & Robbins, C. T. (2006b). Sexual dimorphism, reproductive strategy, and human activities determine resource use by brown bears. Ecology 87: 2636–2646.CrossRefGoogle ScholarPubMed
Rode, K. D., Farley, S. D. & Robbins, C. T. (2006c). Behavioral responses of brown bears mediate nutritional effects of experimentally introduced tourism. Biological Conservation 133: 70–80.CrossRefGoogle Scholar
Rogers, L. L. & Rogers, S. M. (1976). Parasites of bears: a review. International Conference on Bear Research and Management 3: 411–430.Google Scholar
Rush, W. M. (1932). Diphyllobothrium latum in bear. Journal of Mammalogy 13: 274–275.CrossRefGoogle Scholar
Sawaya, M. A., Clevenger, A. P. & Kalinowski, S. T. (2013). Wildlife crossing structures in Banff National Park. Conservation Biology 27: 721–730.CrossRefGoogle ScholarPubMed
Schoen, J. W., Beier, L. R., Lentfer, J. W. & Johnson, L. J. (1987). Denning ecology of brown bears on Admiralty and Chichagof Islands. International Conference on Bear Research and Management 7: 293–304.Google Scholar
Schoen, J. W., Flynn, R. W., Suring, L. H., Titus, K. & Beier, L. R. (1994). Habitat-capability model for brown bear in southeast Alaska. International Conference on Bear Research and Management 9: 317–337.Google Scholar
Schullery, P. (2002). Lewis and Clark among the grizzlies, legend and legacy in the American West. Guildford, CT: The Globe Pequot Press.Google Scholar
Schwartz, C. C., Keating, K. A., Reynolds, H. V. III, et al. (2003a). Reproductive maturation and senescence in the female brown bear. Ursus 14: 109–119.Google Scholar
Schwartz, C. C., Miller, S. D. & Haroldson, M. A. (2003b). Grizzly bear. In: Feldhamer, G. A., Thompson, B. C. & Chapman, J. A. (Eds.), Wild mammals of North America: Biology, management, and conservation (2nd edition, pp. 556–586). Baltimore, MD: Johns Hopkins University Press.Google Scholar
Schwartz, C. C., Haroldson, M. A., White, G. C., et al. (2006). Temporal, spatial, and environmental influences on the demographics of grizzly bears in the Greater Yellowstone Ecosystem. Wildlife Monographs 161: 1–68.CrossRefGoogle Scholar
Schwartz, C. C., Cain, S. L., Podruzny, S. R., Cherry, S. & Frattaroli, L. (2010a). Contrasting activity patterns of sympatric and allopatric black and grizzly bears. Journal of Wildlife Management 74: 1628–1638.CrossRefGoogle Scholar
Schwartz, C. C., Haroldson, M. A. & White, G. C. (2010b). Hazards affecting grizzly bear survival in the greater Yellowstone ecosystem. Journal of Wildlife Management 74: 654–667.CrossRefGoogle Scholar
Schwartz, C. C., Fortin, J. K., Teisberg, J. E., et al. (2014). Body and diet composition of sympatric black and grizzly bears in the Greater Yellowstone Ecosystem. Journal of Wildlife Management 78: 68–78.CrossRefGoogle Scholar
Sellers, R. A. & Aumiller, L. D. (1994). Brown bear population characteristics at McNeil River, Alaska. International Conference on Bear Research and Management 9: 283–293.Google Scholar
Servheen, C. (1983). Grizzly bear food habits, movements and habitat selection in the Mission Mountains, Montana. Journal of Wildlife Management 47: 1026–1035.CrossRefGoogle Scholar
Servheen, C. (1999). Status and management of the grizzly bear in the lower 48 United States. In: Servheen, C., Herrero, S. & Peyton, B. (Eds.), Bears: Status survey and conservation action plan (pp. 50–54). Gland: IUCN/SSC Bear and Polar Bear Specialist Groups, International Union for the Conservation of Nature.Google Scholar
Servheen, C. & Klaver, R. (1983). Grizzly bear dens and denning activity in the Mission and Rattlesnake Mountains, Montana. International Conference on Bear Research and Management 5: 201–207.Google Scholar
Servheen, C., Kasworm, W. & Christensen, A. (1987). Approaches to augmenting grizzly bear populations in the Cabinet Mountains of Montana. International Conference on Bear Research and Management 7: 363–367.Google Scholar
Shields, G. F., Adams, D., Garner, G., et al. (2000). Phylogeography of mitochondrial DNA variation in brown bears and polar bears. Molecular Phylogenetics and Evolution 15: 319–326.CrossRefGoogle ScholarPubMed
Shores, C. R., Mikle, N. & Graves, T. A. (2019). Mapping a keystone shrub species, huckleberry (Vaccinium membranaceum), using seasonal colour change in the Rocky Mountains. International Journal of Remote Sensing 40: 5695–5715.CrossRefGoogle Scholar
Sibly, R. M. & Brown, J. H. (2009). Mammal reproductive strategies driven by offspring mortality–size relationships. American Naturalist 173: E185–E199.CrossRefGoogle ScholarPubMed
Smith, T. S. & Partridge, S. T. (2004). Dynamics of intertidal foraging by coastal brown bears in Southwestern Alaska. Journal of Wildlife Management 68: 233–240.CrossRefGoogle Scholar
Stebler, A. M. (1972). Conservation of the grizzly: ecologic and cultural considerations. International Conference on Bear Research and Management 2: 297–303.Google Scholar
Stenhouse, G. B., Boulanger, J., Lee, J., et al. (2005). Grizzly bear associations along the eastern slopes of Alberta. Ursus 16: 31–40.CrossRefGoogle Scholar
Stewart, B. P., Nelson, T. A., Laberee, K., et al. (2013). Quantifying grizzly bear selection of natural and anthropogenic edges. Journal of Wildlife Management 77: 957–964.CrossRefGoogle Scholar
Steyaert, S. M. J. G., Endrestøl, A., Hackländer, K., Swenson, J. E. & Zedrosser, A. (2012). The mating system of the brown bear Ursus arctos. Mammal Review 42: 12–34.CrossRefGoogle Scholar
Stirling, I. & Derocher, A. E. (1990). Factors affecting the evolution and behavioral ecology of the modern bears. International Conference on Bear Research and Management 8: 189–204.Google Scholar
Stonorov, D. & Stokes, A. W. (1972). Social behavior of the Alaska brown bear. International Conference on Bear Research and Management 2: 232–242.Google Scholar
Storer, T. I. & Tevis, L. P. (1955). California grizzly. Berkeley, CA: University of California Press.Google Scholar
Stringham, S. F. (1990). Grizzly bear reproductive rate relative to body size. International Conference for Bear Research and Management 8: 433–443.Google Scholar
Swenson, J. E. & Haroldson, M. A. (2008). Recent observations of mixed-age litters in brown bears. Ursus 19: 73–79.CrossRefGoogle Scholar
Swenson, J. E., Dahle, B. & Sandegren, F. (2001a). Intraspecific predation in Scandinavian brown bears older than cubs-of-the-year. Ursus 12: 81–92.Google Scholar
Swenson, J. E., Sandegren, F., Brunberg, S. & Segerström, P. (2001b). Factors associated with loss of brown bear cubs in Sweden. Ursus 12: 69–80.Google Scholar
Talbot, S. L. & Shields, G. F. (1996). A phylogeny of the bears (Ursidae) inferred from complete sequences of three mitochondrial genes. Molecular Phylogenetics and Evolution 5: 567–575.CrossRefGoogle ScholarPubMed
Tardiff, S. E. & Stanford, J. A. (1998). Grizzly bear digging: effects on subalpine meadow plants in relation to mineral nitrogen availability. Ecology 79: 2219–2228.CrossRefGoogle Scholar
Troyer, W. A. & Hensel, R. J. (1969). The brown bear of Kodiak Island. Kodiak, AK: US Department of the Interior, Bureau of Sport Fish and Wildlife, Branch of Wildlife Refuges.Google Scholar
US Department of the Interior. (2017). Draft environmental impact statement/grizzly bear restoration plan, North Cascades Ecosystem, Washington. Federal Register 82 FR 4149–4768.Google Scholar
US Fish and Wildlife Service. (1993). Grizzly bear recovery plan. Missoula, MT: US Fish and Wildlife Service, Office of the Grizzly Bear Recovery Coordinator.Google Scholar
US Fish and Wildlife Service. (2000). Record of decision and statement of findings for the Environmental Impact Statement on grizzly bear recovery in the Bitterroot Ecosystem and final rule on establishment of a nonessential experimental population of grizzly bears in the Bitterroot area of Idaho and Montana. Missoula, MT: US Fish and Wildlife Service, Office of the Grizzly Bear Recovery Coordinator.Google Scholar
Van Daele, L. J., Barnes, V. G. & Smith, R. B. (1990). Denning characteristics of brown bears on Kodiak Island, Alaska. International Conference on Bear Research and Management 8: 257–267.Google Scholar
Van Daele, M. B., Robbins, C. T., Semmens, B. X., et al. (2013). Salmon consumption by Kodiak brown bears (Ursus arctos middendorffi) with ecosystem management implications. Canadian Journal of Zoology 91: 164–174.CrossRefGoogle Scholar
van Manen, F. T. & Haroldson, M. A. (2017). Reproduction, survival, and population growth. In: White, P. J., Gunther, K. A. & van Manen, F. T. (Eds.), Yellowstone grizzly bears: Ecology and conservation of an icon of wildness (pp. 29–45). Yellowstone National Park, WY: Yellowstone Forever.Google Scholar
van Manen, F. T., Haroldson, M. A., Bjornlie, D. D., et al. (2016). Density dependence, whitebark pine decline, and changing vital rates of Yellowstone grizzly bears. Journal of Wildlife Management 80: 300–313.CrossRefGoogle Scholar
Vroom, G. W., Herrero, S. & Ogilvie, R. T. (1980). The ecology of winter den sites of grizzly bears in Banff National Park, Alberta. International Conference on Bear Research and Management 4: 321–330.Google Scholar
Waits, L. P., Talbot, S. L., Ward, R. H. & Shields, G. F. (1998). Mitochondrial DNA phylogeography of the North American brown bears and implications for conservation. Conservation Biology 12: 408–417.CrossRefGoogle Scholar
Waits, L. P., Paetkau, D. & Strobeck, C. (1999). Genetics of the bears of the world. In: Servheen, C., Herrero, S. & Peyton, B. (Eds.), Bears: Status survey and conservation action plan (pp. 25–32). Gland: IUCN/SSC Bear and Polar Bear Specialist Groups, International Union for the Conservation of Nature.Google Scholar
Wakkinen, W. L. & Kasworm, W. F. (2004). Demographics and population trends of grizzly bears in the Cabinet-Yaak and Selkirk Ecosystems of British Columbia, Idaho, Montana, and Washington. Ursus 15: 65–75.2.0.CO;2>CrossRefGoogle Scholar
Waller, J. S. & Mace, R. D. (1997). Grizzly bear habitat selection in the Swan Mountains, Montana. Journal of Wildlife Management 61: 1032–1039.CrossRefGoogle Scholar
White, D. Jr., Berardinelli, J. G. & Aune, K. E. (1998a). Reproductive characteristics of the male grizzly bear in the continental United States. Ursus 10: 497–501.Google Scholar
White, D. Jr., Kendall, K. C. & Picton, H. D. (1998b). Grizzly bear feeding activity at alpine army cutworm moth aggregation sites in northwest Montana. Canadian Journal of Zoology 76: 221–227.CrossRefGoogle Scholar
White, D. Jr., Kendall, K. C. & Picton, H. D. (1998c). Seasonal occurrence, body composition, and migration potential of army cutworm moths in northwest Montana. Canadian Journal of Zoology 76: 835–842.CrossRefGoogle Scholar
Wielgus, R. B. & Bunnell, F. L. (1994a). Sexual segregation and female grizzly bear avoidance of males. Journal of Wildlife Management 58: 405–413.CrossRefGoogle Scholar
Wielgus, R. B. & Bunnell, F. L. (1994b). Dynamics of a small, hunted brown bear Ursus arctos population in southwestern Alberta, Canada. Biological Conservation 67: 161–166.CrossRefGoogle Scholar
Wielgus, R. B., Bunnell, F. L., Wakkinen, W. L. & Zager, P. E. (1994). Population dynamics of Selkirk Mountain grizzly bears. Journal of Wildlife Management 58: 266–272.CrossRefGoogle Scholar
Wielgus, R. B., Bunnell, F. L. & Press, A. (1995). Tests of hypotheses for sexual segregation in grizzly bears. Journal of Wildlife Management 59: 552–560.CrossRefGoogle Scholar
Wilk, R. J., Solberg, J. W., Berns, V. D. & Sellers, R. A. (1988). Brown bear (Ursus arctos) with six young. The Canadian Field-Naturalist 102: 541–543.Google Scholar
Wilson, S. M., Madel, M. J., Mattson, D. J., Graham, J. M. & Merrill, T. (2006). Landscape conditions predisposing grizzly bears to conflicts on private agricultural lands in the western USA. Biological Conservation 130: 47–59.CrossRefGoogle Scholar
Wirsing, A. J., Quinn, T. P., Cunningham, C. J., et al. (2018) Alaskan brown bears (Ursus arctos) aggregate and display fidelity to foraging neighborhoods while preying on Pacific salmon along small streams. Ecology and Evolution 8: 9048–9061.CrossRefGoogle ScholarPubMed
Young, D. D. & McCabe, T. R. (1997). Grizzly bear predation rates on caribou calves in northeastern Alaska. Journal of Wildlife Management 61: 1056–1066.CrossRefGoogle Scholar
Zager, P. & Beecham, J. (2006). The role of American black bears and brown bears as predators on ungulates in North America. Ursus 17: 95–109.CrossRefGoogle Scholar
Zarnke, R. L. & Evans, M. B. (1989). Serologic survey for infectious canine hepatitis virus in grizzly bears (Ursus arctos) from Alaska, 1973 to 1987. Journal of Wildlife Diseases 25: 568–573.CrossRefGoogle ScholarPubMed
Zavatsky, B. P. (1976). The use of the skull in age determination of the brown bear. International Conference for Bear Research and Management 4: 275–279.Google Scholar