Carolyn J. Callaghan, Zoology Department, University of Guelph, Guelph, ON N1G 2W1, Canada; Douglas Smith, Yellowstone Center for Resources, PO Box 621, Yellowstone National Park, WY 82190, USA; Paul C. Paquet, Conservation Biology Institute, PO Box 150, Meacham, SK S0K 2V0, Canada; Timmothy Kaminski, Nez Perce Tribe, PO Box 365, Lapwai, ID 83540, USA

Survival is a critical population process and estimating survival rates is an important part of measuring viability of populations. Management of protected wolf (Canis lupus) populations requires quantitative survival measurements so causes of mortality can be identified and unnatural causes reduced. We calculated survival and cause-specific mortality rates for wolves in 7 - 19 packs occupying Banff, Kootenay, Yoho and Yellowstone National Parks for the period 1995 - 1999. Causes of mortality were categorized as highway, railway, shooting or trapping, depredation control, natural accidents, human-caused accident and unknown. Mortality data were obtained from radiocollared and uncollared wolves. Population counts were derived for the late winter period through aerial counts and snow tracking. We further partitioned survival data of wolves into protected and unprotected areas. Protected areas were assigned to wolves whose territory encompassed areas not exposed to roads, rail, harvesting or depredation control pressure; unprotected areas were assigned to collared wolves whose territories were exposed to roads, rail, harvesting or depredation control pressure. For comparison, data were divided into the Canadian National Parks and Yellowstone National Park. Mean annual mortality rate was 32.5% of the Banff, Kootenay and Yoho wolf population under study and 17.5% of the Yellowstone wolf population. Human-caused deaths comprised 73.2% of the total wolf mortalities in the Canadian parks and 44.4% of total wolf mortalities in the Yellowstone area. Annual average human-caused mortalities comprised 73.6% of wolf mortalities in the Canadian parks and 47.3% of wolf mortalities in Yellowstone. Highway and rail collisions were the primary cause of death (29%) in the Canadian parks and unknown or natural deaths were the primary cause of death (55.5%) in Yellowstone. Wolf deaths occurring beyond park boundaries comprised 19.6% of total wolf deaths from the Canadian parks and 28.5% of total wolf deaths from Yellowstone. We discuss implications of wolf mortality and the role of protected areas in the context of population viability.