Food-safety trade-offs drive dynamic behavioural antipredator responses among snowshoe hares.

Prey adopt various antipredator responses to minimize the risk of predation, and the fitness costs of antipredator responses can have emergent effects on the population dynamics of prey species. While the trade-off between food acquisition and predation avoidance has long been recognized in predicting antipredator responses, less attention has been paid to the dynamics of the food-safety trade-off driven by temporal variation in multiple risk factors under changing seasonal conditions. Here, we monitored foraging and vigilance behaviour of a central prey species, snowshoe hares (Lepus americanus), at fine temporal scales over the winter with various types of predation risk, while also experimentally manipulating predation risk by attracting predators to foraging patches. Hares increased foraging and decreased vigilance over the winter, but hares under chronic risk decreased their antipredator efforts to a lesser degree, indicating that those individuals prioritized risk avoidance over food acquisition. Hares also decreased foraging and increased antipredator efforts in response to the temporal activity of predators and environmental cues of predation risk. However, the magnitude of the responses to the environmental cues was mediated by time of winter. While we did not detect a reactive response of hares to acute risk, we did find that hares exhibiting camouflage mismatch proactively increased vigilance. Overall, our results highlight the importance of species-specific traits and changing seasonal conditions in addition to temporal variation in multiple risk factors in predicting antipredator responses and the context dependence of risk effects.

When death comes: linking predator-prey activity patterns to timing of mortality to understand predation risk.

The assumption that activity and foraging are risky for prey underlies many predator-prey theories and has led to the use of predator-prey activity overlap as a proxy of predation risk. However, the simultaneous measures of prey and predator activity along with timing of predation required to test this assumption have not been available. Here, we used accelerometry data on snowshoe hares (Lepus americanus) and Canada lynx (Lynx canadensis) to determine activity patterns of prey and predators and match these to precise timing of predation. Surprisingly we found that lynx kills of hares were as likely to occur during the day when hares were inactive as at night when hares were active. We also found that activity rates of hares were not related to the chance of predation at daily and weekly scales, whereas lynx activity rates positively affected the diel pattern of lynx predation on hares and their weekly kill rates of hares. Our findings suggest that predator-prey diel activity overlap may not always be a good proxy of predation risk, and highlight a need for examining the link between predation and spatio-temporal behaviour of predator and prey to improve our understanding of how predator-prey behavioural interactions drive predation risk.

Phenotypic variation in the molt characteristics of a seasonal coat color-changing species reveals limited resilience to climate change.

The snowshoe hare (Lepus americanus) possesses a broad suite of adaptations to winter, including a seasonal coat color molt. Recently, climate change has been implicated in the range contraction of snowshoe hares along the southern range boundary. With shortening snow season duration, snowshoe hares are experiencing increased camouflage mismatch with their environment reducing survival. Phenological variation of hare molt at regional scales could facilitate local adaptation in the face of climate change, but the level of variation, especially along the southern range boundary, is unknown. Using a network of trail cameras and historical museum specimens, we (1) developed contemporary and historical molt phenology curves in the Upper Great Lakes region, USA, (2) calculated molt rate and variability in and among populations, and (3) quantified the relationship of molt characteristics to environmental conditions for snowshoe hares across North America. We found that snowshoe hares across the region exhibited similar fall and spring molt phenologies, rates and variation. Yet, an insular island population of hares on Isle Royale National Park, MI, completed their molt a week earlier in the fall and initiated molt almost 2 weeks later in the spring as well as exhibited slower rates of molting in the fall season compared to the mainland. Over the last 100 years, snowshoe hares across the region have not shifted in fall molt timing; though contemporary spring molt appears to have advanced by 17 days (~ 4 days per decade) compared to historical molt phenology. Our research indicates that some variation in molt phenology exists for snowshoe hares in the Upper Great Lakes region, but whether this variation is enough to offset the consequences of climate change remains to be seen.

Balancing food acquisition and predation risk drives demographic changes in snowshoe hare population cycles.

Snowshoe hare cycles are one of the most prominent phenomena in ecology. Experimental studies point to predation as the dominant driving factor, but previous experiments combining food supplementation and predator removal produced unexplained multiplicative effects on density. We examined the potential interactive effects of food limitation and predation in causing hare cycles using an individual-based food-supplementation experiment over-winter across three cycle phases that naturally varied in predation risk. Supplementation doubled over-winter survival with the largest effects occurring in the late increase phase. Although the proximate cause of mortality was predation, supplemented hares significantly decreased foraging time and selected for conifer habitat, potentially reducing their predation risk. Supplemented hares also lost less body mass which resulted in the production of larger leverets. Our results establish a mechanistic link between how foraging time, mass loss and predation risk affect survival and reproduction, potentially driving demographic changes associated with hare cycles.

Food availability and long-term predation risk interactively affect antipredator response.

Food availability and temporal variation in predation risk are both important determinants of the magnitude of antipredator responses, but their effects have rarely been examined simultaneously, particularly in wild prey. Here, we determine how food availability and long-term predation risk affect antipredator responses to acute predation risk by monitoring the foraging response of free-ranging snowshoe hares (Lepus americanus) to an encounter with a Canada lynx (Lynx canadensis) in Yukon, Canada, over four winters (2015-2016 to 2018-2019). We examined how this response was influenced by natural variation in long-term predation risk (2-month mortality rate of hares) while providing some individuals with supplemental food. On average, snowshoe hares reduced foraging time up to 10 h after coming into close proximity (≤75 m) with lynx, and reduced foraging time an average of 15.28 ± 7.08 min per lynx encounter. Hares tended to respond more strongly when the distance to lynx was shorter. More importantly, the magnitude of hares' antipredator response to a lynx encounter was affected by the interaction between food-supplementation and long-term predation risk. Food-supplemented hares reduced foraging time more than control hares after a lynx encounter under low long-term risk, but decreased the magnitude of the response as long-term risk increased. In contrast, control hares increased the magnitude of their response as long-term risk increased. Our findings show that food availability and long-term predation risk interactively drive the magnitude of reactive antipredator response to acute predation risk. Determining the factors driving the magnitude of antipredator responses would contribute to a better understanding of the indirect effects of predators on prey populations.

Room without a view-Den excavation in relation to body size in brown bears.

Hibernation is an adaptive strategy to survive harsh winter conditions and food shortage. The use of well-insulated winter dens helps animals minimize energy loss during hibernation. Brown bears (Ursus arctos) commonly use excavated dens for hibernation. Physical attributes of excavated dens are expected to impact the bear's heat retention and energy conservation. The objective of this study was to examine the determinants of cavity size of excavated dens and the impact of physical attributes of excavated dens on energy conservation in hibernating bears, hypothesizing that bears excavate dens in a way to minimize heat loss and optimize energy conservation during hibernation. We predicted that den cavity size would be determined by the bear's body size and that older bears would excavate better-fitting cavities to minimize heat loss, due to their previous experience. We further predicted that physical attributes of excavated dens would affect the bears' posthibernation body condition. Our results revealed that bears excavated a den cavity in relation to their body size, regardless of sex, and that older bears tended to excavate better-fitting den cavities compared to young bears, as we expected. Older bears excavated better-fitting den cavities, suggesting a potentially experience-based shift with age in den-excavation behavior and an optimum cavity size relative to a bear's body size. Our key finding is that insulation of excavated dens provided by wall/rood thickness and bedding materials had a significant positive effect on bears' posthibernation body condition. We believe that our study provides new insight into how not only the quality of denning habitat, but also the quality of dens may affect hibernating animals, by presenting a potential adaptive aspect of den preparation (age effect on efficiency in den excavation) and effect of den attributes on the posthibernation body condition of brown bears.