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Landscape connectivity is essential for maintaining viable populations, particularly for species restricted to fragmented habitats or naturally arrayed in metapopulations and facing rapid climate change. The importance of assessing both structural connectivity (physical distribution of favorable habitat patches) and functional connectivity (how species move among habitat patches) for managing such species is well understood. However, the degree to which functional connectivity for a species varies among landscapes, and the resulting implications for conservation, have rarely been assessed. We used a landscape genetics approach to evaluate resistance to gene flow and, thus, to determine how landscape and climate-related variables influence gene flow for American pikas (Ochotona princeps) in eight federally managed sites in the western United States. We used empirically derived, individual-based landscape resistance models in conjunction with predictive occupancy models to generate patch-based network models describing functional landscape connectivity. Metareplication across landscapes enabled identification of limiting factors for dispersal that would not otherwise have been apparent. Despite the cool microclimates characteristic of pika habitat, south-facing aspects consistently represented higher resistance to movement, supporting the previous hypothesis that exposure to relatively high temperatures may limit dispersal in American pikas. We found that other barriers to dispersal included areas with a high degree of topographic relief, such as cliffs and ravines, as well as streams and distances greater than 1–4 km depending on the site. Using the empirically derived network models of habitat patch connectivity, we identified habitat patches that were likely disproportionately important for maintaining functional connectivity, areas in which habitat appeared fragmented, and locations that could be targeted for management actions to improve functional connectivity. We concluded that climate change, besides influencing patch occupancy as predicted by other studies, may alter landscape resistance for pikas, thereby influencing functional connectivity through multiple pathways simultaneously. Spatial autocorrelation among genotypes varied across study sites and was largest where habitat was most dispersed, suggesting that dispersal distances increased with habitat fragmentation, up to a point. This study demonstrates how landscape features linked to climate can affect functional connectivity for species with naturally fragmented distributions, and reinforces the importance of replicating studies across landscapes.

Although climate acts as a fundamental constraint on the distribution of organisms, understanding how this relationship between climate and distribution varies over a species range is critical for addressing the potential impacts of accelerated climate change on biodiversity. Bioclimatic niche models provide compelling evidence that many species will experience range shifts under scenarios of global change, yet these broad, macroecological perspectives lack specificity at local scales, where unique combinations of environment, biota, and history conspire against generalizations. We explored how these idiosyncrasies of place affect the climate-distribution relationship of the American pika (Ochotona princeps) by replicating intensive field surveys across bioclimatic gradients in eight U.S. national parks. At macroecological scales, the importance of climate as a constraint on pika distribution appears unequivocal; forecasts suggest that the species' range will contract sharply in coming decades. However, the species persists outside of its modeled bioclimatic envelope in many locations, fueling uncertainty and debate over its conservation status. Using a Bayesian hierarchical approach, we modeled variation in local patterns of pika distribution along topographic position, vegetation cover, elevation, temperature, and precipitation gradients in each park landscape. We also accounted for annual turnover in site occupancy probabilities. Topographic position and vegetation cover influenced occurrence in all parks. After accounting for these factors, pika occurrence varied widely among parks along bioclimatic gradients. Precipitation by itself was not a particularly influential predictor. However, measures of heat stress appeared most influential in the driest parks, suggesting an interaction between the strength of climate effects and the position of parks along precipitation gradients. The combination of high elevation, cold temperatures, and high precipitation lowered occurrence probabilities in some parks, suggesting an upper elevational limit for pikas in some environments. Our results demonstrate that the idiosyncrasies of place influence both the nature and strength of the climate-distribution relationship for the American pika. Finegrained, but geographically extensive, studies replicated across multiple landscapes offer insights important to assessing the impacts of climate change that otherwise may be masked at macroecological scales. The hierarchical approach to modeling provides a coherent conceptual and technical framework for gaining these insights.

In 2010, the American pika (Ochotona princeps fenisex) was denied federal protection based on limited evidence of persistence in low‐elevation environments. Studies in nonalpine areas have been limited to relatively few environments, and it is unclear whether patterns observed elsewhere (e.g., Bodie, CA) represent other nonalpine habitats. This study was designed to establish pika presence in a new location, determine distribution within the surveyed area, and evaluate influences of elevation, vegetation, lava complexity, and distance to habitat edge on pika site occupancy. In 2011 and 2012, we conducted surveys for American pika on four distinct subalpine lava flows of Newberry National Volcanic Monument, Oregon, USA. Field surveys were conducted at predetermined locations within lava flows via silent observation and active searching for pika sign. Site habitat characteristics were included as predictors of occupancy in multinomial regression models. Above and belowground temperatures were recorded at a subsample of pika detection sites. Pika were detected in 26% (2011) and 19% (2012) of survey plots. Seventy‐four pika were detected outside survey plot boundaries. Lava complexity was the strongest predictor of pika occurrence, where pika were up to seven times more likely to occur in the most complicated lava formations. Pika were two times more likely to occur with increasing elevation, although they were found at all elevations in the study area. This study expands the known distribution of the species and provides additional evidence for persistence in nonalpine habitats. Results partially support the predictive occupancy model developed for pika at Craters of the Moon National Monument, another lava environment. Characteristics of the lava environment clearly influence pika site occupancy, but habitat variables reported as important in other studies were inconclusive here. Further work is needed to gain a better understanding of the species’ current distribution and ability to persist under future climate conditions. This study documents persistence of a newly discovered population of American pika at elevations below those predicted as optimal for the species. Like other lava environments where pika have been recently documented, lava flows at NNVM appear to be serving as thermal refugia for pika, despite summer temperatures which regularly exceed thermal maxima for the species. It is likely that pika inhabit other low‐elevation lava flows in areas that have never been surveyed.

Climate change vulnerability assessments are an important tool for understanding the threat that climate change poses to species and populations, but do not generally yield insight into the spatial variation in vulnerability throughout a species’ habitat. We demonstrate how to adapt the method of ecological‐niche factor analysis (ENFA) to objectively quantify aspects of species sensitivity to climate change. We then expand ENFA to quantify aspects of exposure and vulnerability to climate change as well, using future projections of global climate models. This approach provides spatially‐explicit insight into geographic patterns of vulnerability, relies only on readily‐available spatial data, is suitable for a wide range of species and habitats, and invites comparison between different species. We apply our methods to a case study of two species of montane mammals, the American pika Ochotona princeps and the yellow‐bellied marmot Marmota flaviventris.

Some of the most compelling examples of ecological responses to climate change are elevational range shifts of individual species, which have been observed throughout the world. A growing body of evidence, however, suggests substantial mediation of simple range shifts due to climate change by other limiting factors. Understanding limiting factors for a species within different contexts, therefore, is critical for predicting responses to climate change. The American pika (Ochotona princeps) is an ideal species for investigating distributions in relation to climate because of their unusual and well-understood natural history as well as observed shifts to higher elevation in parts of their range. We tested three hypotheses for the climatic or habitat characteristics that may limit pika presence and abundance: summer heat, winter snowpack, and forage availability. We performed these tests using an index of pika abundance gathered in a region where environmental influences on pika distribution have not been well-characterized. We estimated relative pika abundance via scat surveys and quantified climatic and habitat characteristics across two North-Central Rocky Mountain Ranges, the Wind River and Bighorn ranges in Wyoming, USA. Pika scat density was highest at mid-elevations and increased linearly with forage availability in both ranges. Scat density also increased with temperatures conducive to forage plant growth, and showed a unimodal relationship with the number of days below -5 degree C, which is modulated by insulating snowpack. Our results provide support for both the forage availability and winter snowpack hypotheses. Especially in montane systems, considering the context-dependent nature of climate effects across regions and elevations as well as interactions between climatic and other critical habitat characteristics, will be essential for predicting future species distributions.

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.

The American pika (Ochotona princeps) is commonly perceived as a species that is at high risk of extinction due to climate change. The purpose of this review is two-fold: to evaluate the claim that climate change is threatening pikas with extinction, and to summarize the conservation status of the American pika. Most American pikas inhabit major cordilleras, such as the Rocky Mountain, Sierra Nevada, and Cascade ranges. Occupancy of potential pika habitat in these ranges is uniformly high and no discernible climate signal has been found that discriminates between the many occupied and relatively few unoccupied sites that have been recently surveyed. Pikas therefore are thriving across most of their range. The story differs in more marginal parts of the species range, primarily across the Great Basin, where a higher percentage of available habitat is unoccupied. A comprehensive review of Great Basin pikas revealed that occupied sites, sites of recent extirpation, and old sites, were regularly found within the same geographic and climatic space as extant sites, and suggested that pikas in the Great Basin tolerated a broader set of habitat and climatic conditions than previously understood. Studies of a small subset of extirpated sites in the Great Basin and in California found that climate variables (most notably measures of hot temperature) were associated more often with extirpated sites than occupied sites. Importantly, upward contraction of the lower elevation boundary also was found at some sites. However, models that incorporated variables other than climate (such as availability of upslope talus habitat) often were better predictors of site persistence. Many extirpations occurred on small habitat patches, which were subject to stochastic extinction, as informed by a long-term pika metapopulation study in Bodie, California. In addition, several sites may have been compromised by cattle grazing or other anthropogenic factors. In contrast, several low, hot sites (Bodie, Mono Craters, Craters of the Moon National Monument and Preserve, Lava Beds National Monument, Columbia River Gorge) retain active pika populations, demonstrating the adaptive capacity and resilience of pikas in response to adverse environmental conditions. Pikas cope with warm temperatures by retreating into cool interstices of their talus habitat and augment their restricted daytime foraging with nocturnal activity. Pikas exhibit significant flexibility in their foraging tactics and are highly selective in their choice of available vegetation. The trait that places pikas at greatest risk from climate change is their poor dispersal capability. Dispersal is more restricted in hotter environments, and isolated low-elevation sites that become extirpated are unlikely to be recolonized in a warming climate. The narrative that American pikas are going extinct appears to be an overreach. Pikas are doing well across most of their range, but there are limited, low-elevation losses that are likely to be permanent in what is currently marginal pika habitat. The resilience of pikas in the face of climate change, and their ability or inability to persist in marginal, hot environments, will continue to contribute to our understanding of the impact of climate change on individual species.

1. Gut microbial communities provide many physiological functions to their hosts, especially in herbivorous animals. We still lack an understanding of how these microbial communities are structured across hosts in nature, especially within a given host species. Studies on laboratory mice have demonstrated that host genetics can influence microbial community structure, but that diet can overwhelm these genetic effects. 2. We aimed to test these ideas in a natural system, the American pika (Ochotona princeps). First, pikas are high-elevation specialists with significant population structure across various mountain ranges in the USA, allowing us to investigate whether similarities in microbial communities match host genetic differences. Additionally, pikas are herbivorous, with some populations exhibiting remarkable dietary plasticity and consuming high levels of moss, which is exceptionally high in fibre and low in protein. This allows us to investigate adaptations to an herbivorous diet, as well as to the especially challenging diet of moss. 3. Here, we inventoried the microbial communities of pika caecal pellets from various populations using 16S rRNA sequencing to investigate structuring of microbial communities across various populations with different natural diets. 4. Microbial communities varied significantly across populations, and differences in microbial community structure were congruent with genetic differences in host population structure, a pattern known as \"phylosymbiosis.\" 5. Several microbial members (Ruminococcus, Prevotella, Oxalobacter and Coprococcus)were detected across all samples, and thus likely represent a \"core microbiome.\" These genera are known to perform a number of services for herbivorous hosts such as fibre fermentation and the degradation of plant defensive compounds, and thus are likely important for herbivory in pikas. Moreover, pikas that feed on moss harboured microbial communities highly enriched in Melainabacteria. This uncultivable candidate phylum has been proposed to ferment fibre for herbivores, and thus may contribute to the ability of some pika populations to consume high amounts of moss. 6. These findings demonstrate that both host genetics and diet can influence the microbial communities of the American pika. These animals may be novel sources of fibre-degrading microbes. Last, we discuss the implications of population-specific microbial communities for conservation efforts in this species.

Aim: The American pika (Ochotona princeps) appears to have experienced climatemediated upslope range contraction in the Great Basin of North America, but this result has not yet been extended to other portions of the pika's range. Our goals were: first, to determine the environmental parameters that most influence current pika distribution within California; second, to infer whether these constraints explain extirpations that have occurred in California; third, to predict future extirpations; and fourth, to advance methods for assessing the degree to which pikas and other climate-sensitive mammals are threatened by climate change. Location: Historical pika record locations in California, USA, spanning four degrees of latitude and longitude, from Mount Shasta to the southern Sierra Nevada. Methods: We identified 67 precise historical pika record locations and surveyed them exhaustively, over multiple years, to determine whether pika populations persist at those sites. We used an information theoretic approach and logistic regression to model current pika occupancy as a function of 16 environmental variables, tested our best-performing model as a predictor of historical occupancy, and then used our model to predict future pika occupancy given anticipated climate change. Results: Pikas no longer occurred at 10 of 67 (15%) historical sites in California. The best predictors of occupancy were average summer temperature and talus habitat area within a 1-km radius. A logistic model fitted to this relationship correctly predicted current occupancy at 94% of sites and correctly hindcasted past occupancy at 93% of sites, suggesting that the model has strong temporal transferability. Depending on the future climate scenario, our model projects that by 2070 pikas will be extirpated from 39% to 88% of these historical sites in California. Main conclusions: Our simple species distribution model for pikas performs remarkably well for both current and historical periods. Pika distribution appears to be governed primarily by behavioural restrictions mediated by summer temperature and by the configuration of talus habitat available to pikas locally. Pikas, and other montane species in the western USA, may be subjected to above-average exposure to climate change because summer temperature is projected to rise more than annual temperature.

Many organisms leave evidence of their former occurrence, such as scat, abandoned burrows, middens, ancient eDNA or fossils, which indicate areas from which a species has since disappeared. However, combining this evidence with contemporary occurrences within a single modeling framework remains challenging. Traditional binary species‐distribution modeling reduces occurrence to two temporally coarse states (present/absent), so thus cannot leverage the information inherent in temporal sequences of evidence of past occurrence. In contrast, ordinal modeling can use the natural time‐varying order of states (e.g. never occupied versus previously occupied versus currently occupied) to provide greater insights into range shifts. We demonstrate the power of ordinal modeling for identifying the major influences of biogeographic and climatic variables on current and past occupancy of the American pika Ochotona princeps, a climate‐sensitive mammal. Sampling over five years across the species' southernmost, warm‐edge range limit, we tested the effects of these variables at 570 habitat patches where occurrence was classified either as binary or ordinal. The two analyses produced different top models and predictors – ordinal modeling highlighted chronic cold as the most‐important predictor of occurrence, whereas binary modeling indicated primacy of average summer‐long temperatures. Colder wintertime temperatures were associated in ordinal models with higher likelihood of occurrence, which we hypothesize reflect longer retention of insulative and meltwater‐provisioning snowpacks. Our binary results mirrored those of other past pika investigations employing binary analysis, wherein warmer temperatures decrease likelihood of occurrence. Because both ordinal‐ and binary‐analysis top models included climatic and biogeographic factors, results constitute important considerations for climate‐adaptation planning. Cross‐time evidences of species occurrences remain underutilized for assessing responses to climate change. Compared to multi‐state occupancy modeling, which presumes all states occur in the same time period, ordinal models enable use of historical evidence of species' occurrence to identify factors driving species' distributions more finely across time.