Den site selection by male brown bears at the population's expansion front

Brown bears (Ursus arctos) spend about half of the year in winter dens. In order to preserve energy, bears may select denning locations that minimize temperature loss and human disturbance. In expanding animal populations, demographic structure and individual behavior at the expansion front can differ from core areas. We conducted a non-invasive study of male brown bear den sites at the male-biased, low-density western expansion front of the Scandinavian brown bear population, comparing den locations to the available habitat. Compared to the higher-density population core in which intraspecific avoidance may affect den site selection of subordinate bears, we expected resource competition in the periphery to be low, and all bears to be able to select optimal den sites. In addition, bears in the periphery had access to free-ranging domestic sheep during summer. We found that males in the periphery denned on high-elevation slopes, probably providing good drainage, longer periods of consistent, insulating snow cover and fewer melting-freezing events. Forests were the principal denning habitat and no dens were found in alpine areas. The Scandinavian brown bears have a history of intense harvest, including culling at the den. This may have exerted a selection pressure to avoid denning in open alpine habitat which compared to forests provide little cover. The bears denned away from main roads and in steep, rugged terrain, probably limiting human access. The odds for finding a bear den decreased with increasing distance to the population core where females could be found. Previous studies have documented directed movement of male brown bears from the male-biased population periphery toward the core areas during the mating season. In this way, denning males may be trading off between low resource competition and access to sheep in the low-density periphery, and mating opportunities in the higher-density population core.