The influence of winter severity, predation and senescence on moose habitat use.

Habitat use is widely known to be influenced by abiotic and biotic factors, such as climate, population density, foraging opportunity and predation risk. The influence of the life-history state of an individual organism on habitat use is less well understood, especially for terrestrial mammals. There is good reason to expect that life-history state would affect habitat use. For example, organisms exhibiting poor condition associated with senescence have an increased vulnerability to predation and that vulnerability is known to alter habitat use strategies. We assessed the influence of life-history stage on habitat use for 732 moose (Alces alces) killed by wolves (Canis lupus) over a 50-year period in Isle Royale National Park, an island ecosystem in Lake Superior, USA. We developed regression models to assess how location of death was associated with a moose's life-history stage (prime-aged or senescent), presence or absence of senescent-associated pathology (osteoarthritis and jaw necrosis), and annual variation in winter severity, moose density and ratio of moose to wolves, which is an index of predation risk. Compared to senescent moose, prime-aged moose tend to make greater use of habitat farther from the shoreline of Isle Royale. That result is ecologically relevant because shoreline habitat on Isle Royale tends to provide better foraging opportunities for moose but is also associated with increased predation risk. During severe winters prime-aged moose tend to make greater use of habitat that is closer to shore in relation to senescent-aged moose. Furthermore, moose of both age classes were more likely to die in riskier, shoreline habitat during years when predation risk was lower in the preceding year. Our results highlight a complicated connection between life history, age-structured population dynamics and habitat-related behaviour. Our analysis also illustrates why intraspecific competition should not be the presumed mechanism underlying density-dependent habitat use, if predation risk is related to density, as it is expected to be in many systems.