Ecological theory predicts that the diffuse risk cues generated by wide-ranging, active predators should induce prey behavioural responses but not major, population- or community-level consequences. We evaluated the non-consumptive effects (NCEs) of an active predator, the grey wolf (Canis lupus), by simultaneously tracking wolves and the behaviour, body fat, and pregnancy of elk (Cervus elaphus), their primary prey in the Greater Yellowstone Ecosystem. When wolves approached within 1 km, elk increased their rates of movement, displacement and vigilance. Even in high-risk areas, however, these encounters occurred only once every 9 days. Ultimately, despite 20-fold variation in the frequency of encounters between wolves and individual elk, the risk of predation was not associated with elk body fat or pregnancy. Our findings suggest that the ecological consequences of actively hunting large carnivores, such as the wolf, are more likely transmitted by consumptive effects on prey survival than NCEs on prey behaviour.

Conservation of migration requires information on behavior and environmental determinants. The spatial distribution of forage resources, which migration exploits, often are altered and may have subtle, unintended consequences. Supplemental feeding is a common management practice, particularly for ungulates in North America and Europe, and carryover effects on behavior of this anthropogenic manipulation of forage are expected in theory, but have received limited empirical evaluation, particularly regarding effects on migration. We used global positioning system (GPS) data to evaluate the influence of winter feeding on migration behavior of 219 adult female elk (Cervus elaphus) from 18 fed ranges and 4 unfed ranges in western Wyoming. Principal component analysis revealed that the migratory behavior of fed and unfed elk differed in distance migrated, and the timing of arrival to, duration on, and departure from summer range. Fed elk migrated 19.2 km less, spent 11 more days on stopover sites, arrived to summer range 5 days later, resided on summer range 26 fewer days, and departed in the autumn 10 days earlier than unfed elk. Time-to-event models indicated that differences in migratory behavior between fed and unfed elk were caused by altered sensitivity to the environmental drivers of migration. In spring, unfed elk migrated following plant green-up closely, whereas fed elk departed the feedground but lingered on transitional range, thereby delaying their arrival to summer range. In autumn, fed elk were more responsive to low temperatures and precipitation events, causing earlier departure from summer range than unfed elk. Overall, supplemental feeding disconnected migration by fed elk from spring green-up and decreased time spent on summer range, thereby reducing access to quality forage. Our findings suggest that ungulate migration can be substantially altered by changes to the spatial distribution of resources, including those of anthropogenic origin, and that management practices applied in one season may have unintended behavioral consequences in subsequent seasons.

Polar bears (Ursus maritimus) summer on the sea ice or, where it melts, on shore. Although the physiology of "ice" bears in summer is unknown, "shore" bears purportedly minimize energy losses by entering a hibernation-like state when deprived of food. Such a strategy could partially compensate for the loss of on-ice foraging opportunities caused by climate change. However, here we report gradual, moderate declines in activity and body temperature of both shore and ice bears in summer, resembling energy expenditures typical of fasting, nonhibernating mammals. Also, we found that to avoid unsustainable heat loss while swimming, bears employed unusual heterothermy of the body core. Thus, although well adapted to seasonal ice melt, polar bears appear susceptible to deleterious declines in body condition during the lengthening period of summer food deprivation.

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