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Temporal variation in stress might signify changes in an animal’s internal or external environment, while spatial variation in stress might signify variation in the quality of the habitats that individual animals experience. Habitat-induced variations in stress might be easiest to detect in highly territorial animals, and especially in species that do not take advantage of common strategies for modulating habitat-induced stress, such as migration (escape in space) or hibernation (escape in time). Spatial and temporal variation in response to potential stressors has received little study in wild animals, especially at scales appropriate for relating stress to specific habitat characteristics. Here, we use the American pika (Ochotona princeps), a territorial small mammal, to investigate stress response within and among territories. For individually territorial animals such as pikas, differences in habitat quality should lead to differences in stress exhibited by territory owners. We indexed stress using stress-associated hormone metabolites in feces collected non-invasively from pika territories every 2 weeks from June to September 2018. We hypothesized that differences in territory quality would lead to spatial differences in mean stress and that seasonal variation in physiology or the physical environment would lead to synchronous variation across territories through time. We used linear mixed-effects models to explore spatiotemporal variation in stress using fixed effects of day-of-year and broad habitat characteristics (elevation, aspect, site), as well as local variation in habitat characteristics hypothesized to affect territory quality for this saxicolous species (talus depth, clast size, available forage types). We found that temporal variation within territories was greater than spatial variation among territories, suggesting that shared seasonal stressors are more influential than differences in individual habitat quality. This approach could be used in other wildlife studies to refine our understanding of habitat quality and its effect on individual stress levels as a driver of population decline.

Behaviour is an important mechanism for accommodating rapid environmental changes. Understanding a species' capacity for behavioural plasticity is therefore a key, but understudied, aspect of developing tractable conservation and management plans under climate-change scenarios. Here, we quantified behavioural differences between American pikas (Ochotona princeps) living in an atypical, low-elevation habitat versus those living in a more-typical, alpine habitat. With respect to foraging strategy, low-elevation pikas spent more time consuming vegetation and less time caching food for winter, compared to high-elevation pikas. Low-elevation pikas were also far more likely to be detected in forested microhabitats off the talus than their high-elevation counterparts at midday. Finally, pikas living in the atypical habitat had smaller home range sizes compared to those in typical habitat or any previously published home ranges for this species. Our findings indicate that behavioural plasticity likely allows pikas to accommodate atypical conditions in this low-elevation habitat, and that they may rely on critical habitat factors such as suitable microclimate refugia to behaviourally thermoregulate. Together, these results suggest that behavioural adjustments are one important mechanism by which pikas can persist outside of their previously appreciated dietary and thermal niches.

Range contractions in the Great Basin over the last century suggest that American pikas (Ochotona princeps) might be highly sensitive to climate change. However, documentation of pikas residing at relatively warm, low-elevation sites has recently shed new light on the possible resilience of pika populations to warmer ambient conditions when they have access to cooler microhabitats for thermoregulation. To provide insight into the possible behavioral mechanisms of adaptation to warmer habitats, we investigated activity patterns, foraging behavior, and space use of a population of pikas living in an atypical, warmer habitat at Craters of the Moon National Monument and Preserve, an extensive lava flow surrounded by high desert grassland and sagebrush communities in southern Idaho. We compared pika behavior at this site to that of a population in a more typical alpine habitat at Grays Peak in the nearby Pioneer Mountains in Idaho. Specifically, we evaluated and compared activity patterns and foraging behavior of pikas in relation to temperature, time of day, and month. Pikas at Craters of the Moon were less active aboveground during all times of the day from late May until mid-August, compared to those at Grays Peak. Surface temperatures were warmer across the entire season at Craters of the Moon than at Grays Peak, and pikas exhibited a crepuscular activity pattern that coincided with variations in surface temperature at Craters of the Moon. Pikas were the least active during the midday, when the surface temperatures were highest and the insulating effect of the lava tubes (i.e., the difference between surface and crevice temperatures) was most pronounced. Pikas at Craters of the Moon spent less time haying and displayed fewer territorial behaviors than pikas at Grays Peak, but both groups filled a similar number of hay piles. The vegetation community was less diverse and sparser at Craters of the Moon than at Grays Peak, and consequently, the vegetation that was consumed and cached reflected these differences. Our results expand the body of literature about American pikas at their environmental limits, and this study is the first step in identifying the unique suite of behaviors that pikas use to persist in a seemingly inhospitable environment at Craters of the Moon.

Mammalian habitat specialists are suffering notable population declines and localized extinctions in response to climate change. Plastic behavioral responses, especially in foraging, may be critical for specialists to tolerate changes in temperature, precipitation, and resource availability. Here, we investigate the foraging behavior of a mammalian alpine specialist living in atypical habitat. American pikas (Ochotona princeps) are typically limited to high elevations in western North America; however, they persist near sea level in the Columbia River Gorge, well outside their previously assumed climatic niche. We hypothesized that utilizing unusual food resources contributes to pika persistence in this unusual climate. Moss comprised more than 60% of the diet at 2 sites, more than observed for any mammalian herbivore in the wild. Moss is available year-round in this habitat; thus, by specializing on moss, pikas do not have to construct large food caches to survive winter. These results suggest a larger degree of behavioral and dietary plasticity than previously assumed for this species. Understanding a species’ capacity to adapt its foraging strategies to new resource landscapes will be essential to assessing its vulnerability to future climate change and to developing conservation plans.

The American pika (Ochotona princeps) is considered a sentinel species for detecting ecological effects of climate change. Pikas are declining within a large portion of their range, and ongoing research suggests loss of sub-surface ice as a mechanism. However, no studies have demonstrated physiological responses of pikas to sub-surface ice features. Here we present the first analysis of physiological stress in pikas living in and adjacent to habitats underlain by ice. Fresh fecal samples were collected non-invasively from two adjacent sites in the Rocky Mountains (one with sub-surface ice and one without) and analyzed for glucocorticoid metabolites (GCM). We also measured sub-surface microclimates in each habitat. Results indicate lower GCM concentration in sites with sub-surface ice, suggesting that pikas are less stressed in favorable microclimates resulting from sub-surface ice features. GCM response was well predicted by habitat characteristics associated with sub-surface ice features, such as lower mean summer temperatures. These results suggest that pikas inhabiting areas without sub-surface ice features are experiencing higher levels of physiological stress and may be more susceptible to changing climates. Although post-deposition environmental effects can confound analyses based on fecal GCM, we found no evidence for such effects in this study. Sub-surface ice features are key to water cycling and storage and will likely represent an increasingly important component of water resources in a warming climate. Fecal samples collected from additional watersheds as part of current pika monitoring programs could be used to further characterize relationships between pika stress and sub-surface ice features.

Wildfires are increasing in frequency and severity as a result of climate change in many ecosystems; however, effects of altered disturbance regimes on wildlife remain poorly quantified. Here, we leverage an unexpected opportunity to investigate how fire affects the occupancy and abundance of a climate-sensitive habitat specialist, the American pika (Ochotona princeps). We determine the effects of a fire on microclimates within talus and explore habitat factors promoting persistence and abundance in fire-affected habitat. During the fire, temperatures in talus interstices remained below 19°C, suggesting that animals could have survived in situ. Within 2 years, pikas were widely distributed throughout burned areas and did not appear to be physiologically stressed at severely burned sites. Furthermore, pika densities were better predicted by topographic variables known to affect this species than by metrics of fire severity. This widespread distribution may reflect quick vegetation recovery and the fact that the fire did not alter the talus microclimates in the following years. Together, these results highlight the value of talus as a thermal refuge for small animals during and after fire. They also underscore the importance of further study in individual species’ responses to typical and altered disturbance regimes.

Predicting habitat suitability under climate change is vital to conserving biodiversity. However, current species distribution models rely on coarse scale climate data, whereas fine scale microclimate data may be necessary to assess habitat suitability and generate predictive models. Here, we evaluate disparities between temperature data at the coarse scale from weather stations versus fine-scale data measured in microhabitats required for a climate-sensitive mammal, the American pika (Ochotona princeps). We collected two years of temperature data in occupied talus habitats predicted to be suitable (high elevation) and unsuitable (low elevation) by the bioclimatic envelope approach. At low elevations, talus surface and interstitial microclimates drastically differed from ambient temperatures measured on-site and at a nearby weather station. Interstitial talus temperatures were frequently decoupled from high ambient temperatures, resulting in instantaneous disparities of over 30 °C between these two measurements. Microhabitat temperatures were also highly heterogeneous, such that temperature measurements within the same patch of talus were not more correlated than measurements at distant patches. An experimental manipulation revealed that vegetation cover may cool the talus surface by up to 10 °C during the summer, which may contribute to this spatial heterogeneity. Finally, low elevation microclimates were milder and less variable than typical alpine habitat, suggesting that, counter to species distribution model predictions, these seemingly unsuitable habitats may actually be better refugia for this species under climate change. These results highlight the importance of fine-scale microhabitat data in habitat assessments and underscore the notion that some critical refugia may be counterintuitive.