Scales of movement by elk (Cervus elaphus) in response to heterogeneity in forage resources and predation risk

Animals may respond to spatial and temporal heterogeneity by altering their movement patterns. The time an animal spends in an area of a given size is termed 'first-passage time' and can be used to identify the scales at which different movement processes occur. Using first-passage time and 2-h observations, we identified nested spatial scales representing three movement behaviours for elk (Cervus elaphus) - inactive/ resting (moves < 50 m), active/foraging (<(x)over bar> = 276.7 m, SD = 56.6), and active/relocating ((x) over bar = 1628.3 m, SD = 436.6). Our ability to identify inactive behaviour was limited by GPS accuracy. The scale separating relocating and foraging behaviour. ranged 550-1650 m across individuals and varied quadratically with the mean patch size of cutover forest in an animal's home range. We classified path segments into the 3 movement behaviours and related behaviours to local. environmental conditions. Elk were likely to be inactive in areas having a low predicted use by wolves (Canis lupus), farther than 50 in from anthropogenic linear clearings, and where microclimatic conditions were cool (high shrub cover and north to east-facing slopes). In contrast, elk were most likely to forage in areas having intermediate levels of herbaceous biomass and low movement costs. Elk were most likely to be relocating when in areas of high wolf use, when close to linear clearings, and in energetically costly situations such as moving upslope. We discuss how elk use of potential foraging habitats may be restricted in this landscape by risks imposed by predators, humans, or both.