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As the climate warms, endotherms are challenged with maintaining body temperature within a neutral range. Modifying behaviour to mitigate heat loads is a potentially low-cost response to avoid heat stress and may be critical to persistence in a changing environment, especially for large endotherms. We tested the hypothesis that bed sites are a thermal refuge and means of behavioural mitigation of heat loads for a large temperate ungulate especially prone to heat stress, the Shiras moose, Alces alces shirasi. We predicted that if bed sites are a thermal refuge, selection of bed sites should be consistent with sites that reduce heat gain or increase heat loss via convection or conduction. Moose selected cool bed sites with a substrate that was moist, wet or had standing water, and avoided bedding in open or sparse vegetation. On warm days and during the warmest periods of the day, moose increased selection for wet substrate and for bed sites in standing water with sparse vegetation, which they avoided on cool days and during cool periods of the day. Strong and context-dependent selection for bed sites that would reduce heat gain and ameliorate heat load supports the notion that bed sites are a thermal refuge and a way to behaviourally mitigate warming temperatures. Flexibility in habitat selection may be a primary way for large endotherms to combat a warming climate, although doing so is contingent on habitat assemblages that offer thermal refuge to limit heat gain and importantly, facilitate heat loss.

Climate warming creates energetic challenges for endothermic species by increasing metabolic and hydric costs of thermoregulation. Although endotherms can invoke an array of behavioral and physiological strategies for maintaining homeostasis, the relative effectiveness of those strategies in a climate that is becoming both warmer and drier is not well understood. In accordance with the heat dissipation limit theory which suggests that allocation of energy to growth and reproduction by endotherms is constrained by the ability to dissipate heat, we expected that patterns of habitat use by large, heat-sensitive mammals across multiple scales are critical for behavioral thermoregulation during periods of potential heat stress and that they must invest a large portion of time to maintain heat balance. To test our predictions, we evaluated mechanisms underpinning the effectiveness of bed sites for ameliorating daytime heat loads and potential heat stress across the landscape while accounting for other factors known to affect behavior. We integrated detailed data on microclimate and animal attributes of moose Alces alces, into a biophysical model to quantify costs of thermoregulation at fine and coarse spatial scales. During summer, moose spent an average of 67.8% of daylight hours bedded, and selected bed sites and home ranges that reduced risk of experiencing heat stress. For most of the day, shade could effectively mitigate the risk of experiencing heat stress up to 10°C, but at warmer temperatures (up to 20°C) wet soil was necessary to maintain homeostasis via conductive heat loss. Consistent selection across spatial scales for locations that reduced heat load underscores the importance of the thermal environment as a driver of behavior in this heat-sensitive mammal. Moose in North America have long been characterized as riparian-obligate species because of their dependence on woody plant species for food. Nevertheless, the importance of dissipating endogenous heat loads conductively through wet soil suggests riparian habitats also are critical thermal refuges for moose. Such refuges may be especially important in the face of a warming climate in which both high environmental temperatures and drier conditions will likely exacerbate limits to heat dissipation, especially for large, heat-sensitive animals.

Accounting for within-species variability in the relationship between occurrence and climate is essential to forecasting species’ responses to climate change. Few climate-vulnerability assessments explicitly consider intraspecific variation, and those that do typically assume that variability is best explained by genetic affinity. Here, we evaluate how well heterogeneity in responses to climate by a cold-adapted mammal, the American pika (Ochotona princeps), aligns with subdivisions of the geographic range by phylogenetic lineage, physiography, elevation or ecoregion. We find that variability in climate responses is most consistently explained by an ecoregional subdivision paired with background sites selected from a broad spatial extent indicative of long-term (millennial-scale) responses to climate. Our work challenges the common assumption that intraspecific variation in climate responses aligns with genetic affinity. Accounting for the appropriate context and scale of heterogeneity in species’ responses to climate will be critical for informing climate-adaptation management strategies at the local (spatial) extents at which such actions are typically implemented. Intraspecies response to climate change is expected to align with genetic affinity. Using the American pika as a case study suggests that divisions of species distributions best explain intraspecific heterogeneity in climate relationships.

Contemporary climate change is altering temperature profiles across the globe. Increasing temperatures can reduce the amount of time during which conditions are suitable for animals to engage in essential activities, such as securing food. Behavioural plasticity, the ability to alter behaviour in response to the environment, may provide animals with a tool to adjust to changes in the availability of suitable thermal conditions. The extent to which individuals can alter fitness-enhancing behaviours, such as food collection, to proximately buffer variation in temperature, however, remains unclear. Even less well understood are the potential performance advantages of flexible strategies among endotherms. We examined the degree to which individuals altered rates of food collection in response to temperature, and two potential benefits, using the American pika (Ochotona princeps), a temperature-sensitive, food-hoarding mammal, as a model. From July-September 2013-2015, we used motion-activated cameras and in situ temperature loggers to examine pika food-caching activity for 72 individuals across 10 sites in the central Rocky Mountains, USA. We quantified % nitrogen by cache volume as a metric of cache quality, and the number of events during which pikas were active in temperatures ≥25°C as a measure of potential thermoregulatory stress. We found a strong negative effect of temperature on the rate at which pikas cached food. Individual responses to temperature varied substantially in both the level of food-collecting activity and in the degree to which individuals shifted activity with warming temperature. After accounting for available foraging time, individuals that exhibited greater plasticity collected a comparable amount of nitrogen, while simultaneously experiencing fewer occasions in which temperatures eclipsed estimated thermal tolerances. By varying food-collection norms of reaction, individuals were able to plastically respond to temperature-driven reductions in foraging time. Through this increased flexibility, individuals amassed food caches of comparable quality, while minimizing exposure to potentially stressful thermal conditions. Our results suggest that, given sufficient resource quality and availability, plasticity in foraging activity may help temperature-limited endotherms adjust to climate-related constraints on foraging time.

Of the primary responses to contemporary climate change – “move, adapt, acclimate, or die” – that are available to organisms, “acclimate” may be effectively achieved through behavioral modification. Behavioral flexibility allows animals to rapidly cope with changing environmental conditions, and behavior represents an important component of a species’ adaptive capacity in the face of climate change. However, there is currently a lack of knowledge about the limits or constraints on behavioral responses to changing conditions. Here, we characterize the contexts in which organisms respond to climate variability through behavior. First, we quantify patterns in behavioral responses across taxa with respect to timescales, climatic stimuli, life-history traits, and ecology. Next, we identify existing knowledge gaps, research biases, and other challenges. Finally, we discuss how conservation practitioners and resource managers can incorporate an improved understanding of behavioral flexibility into natural resource management and policy decisions.

Contemporary climate change affects nearly all biomes, causing shifts in animal distributions and resource availability. Changes in resource selection may allow individuals to offset climatic stress, thereby providing a mechanism for persistence amidst warming conditions. Whereas the role of predation risk in food choice has been studied broadly, the extent to which individuals respond to thermoregulatory risk by changing resource preferences is unclear. We addressed whether individuals compensated for temperature-related reductions in foraging time by altering forage preferences, using the American pika (Ochotona princeps) as a model species. We tested two hypotheses: (1) food-quality hypothesis—individuals exposed to temperature extremes should select higher-quality vegetation in return for accepting a physiologically riskier feeding situation; and (2) food-availability hypothesis—individuals exposed to temperature extremes should prioritize foraging quickly, thereby decreasing selection for higher-quality food. We quantified the composition and quality (% moisture, % nitrogen, and fiber content) of available and harvested vegetation, and deployed a network of temperature sensors to measure in situ conditions for 30 individuals, during July–Sept., 2015. Individuals exposed to more extreme daytime temperatures showed increased selection for high-nitrogen and for low-fiber vegetation, demonstrating strong support for the food-quality hypothesis. By contrast, pikas that experienced warmer conditions did not reduce selection for any of the three vegetation-quality metrics, as predicted by the food-availability hypothesis. By shifting resource-selection patterns, temperature-limited animals may be able to proximately buffer some of the negative effects associated with rapidly warming environments, provided that sufficient resources remain on the landscape.

Contemporary climate change is affecting nearly all biomes, causing shifts in animal distributions, phenology, and persistence. Favorable microclimates may buffer organisms against rapid changes in climate, thereby allowing time for populations to adapt. The degree to which microclimates facilitate the local persistence of climate-sensitive species, however, is largely an open question. We addressed the importance of microrefuges in mammalian thermal specialists, using the American pika (Ochotona princeps) as a model organism. Pikas are sensitive to ambient temperatures, and are active year-round in the alpine where conditions are highly variable. We tested four hypotheses about the relationship between microrefuges and pika occurrence: 1) Local-habitat Hypothesis (local-habitat conditions are paramount, regardless of microrefuge); 2) Surface-temperature Hypothesis (surrounding temperatures, unmoderated by microrefuge, best predict occurrence); 3) Interstitial-temperature Hypothesis (temperatures within microrefuges best predict occurrence), and 4) Microrefuge Hypothesis (the degree to which microrefuges moderate the surrounding temperature facilitates occurrence, regardless of other habitat characteristics). We examined pika occurrence at 146 sites across an elevational gradient. We quantified pika presence, physiographic habitat characteristics and forage availability at each site, and deployed paired temperature loggers at a subset of sites to measure surface and subterranean temperatures. We found strong support for the Microrefuge Hypothesis. Pikas were more likely to occur at sites where the subsurface environment substantially moderated surface temperatures, especially during the warm season. Microrefugium was the strongest predictor of pika occurrence, independent of other critical habitat characteristics, such as forage availability. By modulating surface temperatures, microrefuges may strongly influence where temperature-limited animals persist in rapidly warming environments. As climate change continues to manifest, efforts to understand the changing dynamics of animal-habitat relationships will be enhanced by considering the quality of microrefuges.