Ochotona Princeps Research Articles

Evidence of Intraspecific Adaptive Variation in the American Pika (Ochotona princeps) on a Continental Scale Using a Target Enrichment and Mitochondrial Genome Skimming Approach.

Montane landscapes present an array of abiotic challenges that drive adaptive evolution amongst organisms. These adaptations can promote habitat specialisation, which may heighten the risk of extirpation from environmental change. For example, higher metabolic rates in an endothermic species may contribute to heightened cold tolerance, whilst simultaneously limiting heat tolerance. Here, using the climate-sensitive American pika (Ochotona princeps), we test for evidence of intraspecific adaptive variation amongst environmental gradients across the Intermountain West of North America. We leveraged results from previous studies on pika adaptation to generate a custom nuclear target enrichment design to sequence several hundred candidate genes related to cold, hypoxia and dietary detoxification. We also applied a 'genome skimming' approach to sequence mitochondrial DNA. Using genotype-environment association tests, we identified rare genomic variants associated with elevation and temperature variation amongst populations. Amongst mitochondrial genes, we identified intraspecific variation in selective signals and significant changes to the amino acid property equilibrium constant, which may relate to electron transport chain efficiency. These results illustrate a complex dynamic of adaptive variation amongst O. princeps where lineages and populations have adapted to unique regional conditions. Some of the clearest signals of selection were in a genetic lineage that includes pikas of the Great Basin region, which is also where recent localised extirpations have taken place and highlights the risk of losing adaptive alleles during environmental change.

Fifty-year population trajectory in a marginal American Pika (Ochotona princeps) population

Abstract The American Pika (Ochotona princeps) has been considered a species at risk due to warming temperatures associated with climate change. Many life-history attributes of pikas contribute to the sensitivity of pikas to warming temperatures. Repeated censuses of a marginal (warm, low-elevation) population of pikas at Bodie State Historic Park, California, conducted from 1972 to 2022, are presented to track the population trajectory of pikas for a time period predating recent awareness of global warming to the present day, thus giving a comprehensive portrayal of how American pikas may be responding to climate change. The northern constellation of anthropogenic habitat patches (mine ore dumps) showed no decline in percent of patches occupied or in total number of pika territories over time, suggesting that pikas in that area have not been at risk of extirpation resulting from climate change. In contrast, the pika population in the southern constellation of patches showed significant declines in percent of patches occupied and number of pika territories occupied. That area was unoccupied for about a decade beginning in 2006, but was recently recolonized from the northern constellation of patches. The most likely cause of the initial decline and transient extirpation in the south appears to result from fragmentation and stochastic population dynamics, independent of climatic factors that we investigated. Assessments of climatic impacts on American pikas should take into account the dynamics documented in the Bodie pika population and its resilience over time.

Assessing the Impact of Climate Change on Global Biodiversity: Trends and Predictions

Global warming which is as a result of human activities such as burning of fossil fuels and deforestation is a major threat to species diversity. This study is based on species in the climate change hotspots, using the SSDM, phenology, species traits, responses of ectotherms, and vegetation activity. We evaluate shifts in the distribution and numbers of species and their probabilities of dying out. An average Global warming of 1.1°C since the end of the nineteenth century has altered precipitation, deteriorated habitats, and increased the rate of species’ disappearance. If emissions are not reduced, 20-30% of species like the American Pika and many amphibians could be wiped out by end of the century. Phenological activities are taking place 2.5 days earlier per decade, which affects the balance of ecosystems. This study also stresses the importance of flexibility in conservation approaches like connecting the fragmented habitats, rehabilitating the disturbed ones, and the protection of ecological connectivity to reduce the impacts of climate change on species.

Retrospective review of the pathology of American pikas.

American pikas (Ochotona princeps) are small lagomorphs that live in mountainous talus areas of western North America. Studies on the histopathology of American pikas are limited. We summarize here the clinical histories, and gross and histologic findings of 12 American pikas, including 9 captive (wild-caught) and 3 wild animals. Death was often attributed to stress (transport, handling, anesthesia) with few-to-no premonitory clinical signs. Infection was the cause of death in 2 cases: 1 had bacterial pyogranulomatous dermatitis, cellulitis, and lymphadenitis with sepsis; the other case had oomycete-induced necrotizing colitis. Incidental parasitic infections included sarcocystosis, nematodosis (oxyurids), and ectoparasitism. Most animals with adequate nutritional status had periportal hepatic lipidosis; this finding was absent in all animals with adipose atrophy, and it is possible that periportal hepatic lipidosis is non-pathologic in American pikas. Three cases had myocardial necrosis that was considered the cause of death; the cause of necrosis was not determined, but it may have been caused by stress or vitamin E-selenium deficiency. Esophageal hyperkeratosis was noted in animals with a history of anorexia and negative energy balance; accumulation of esophageal keratin can result from lack of mucosal abrasion by ingesta. Several histologic findings that are likely normal in American pikas include splenic extramedullary hematopoiesis, thymic tissue in adults, and Clostridium sp. in the enteric lumen.

Putative climate adaptation in American pikas (Ochotona princeps) is associated with copy number variation across environmental gradients

Improved understanding of the genetic basis of adaptation to climate change is necessary for maintaining global biodiversity moving forward. Studies to date have largely focused on sequence variation, yet there is growing evidence that suggests that changes in genome structure may be an even more significant source of adaptive potential. The American pika (Ochotona princeps) is an alpine specialist that shows some evidence of adaptation to climate along elevational gradients, but previous work has been limited to single nucleotide polymorphism based analyses within a fraction of the species range. Here, we investigated the role of copy number variation underlying patterns of local adaptation in the American pika using genome-wide data previously collected across the entire species range. We identified 37–193 putative copy number variants (CNVs) associated with environmental variation (temperature, precipitation, solar radiation) within each of the six major American pika lineages, with patterns of divergence largely following elevational and latitudinal gradients. Genes associated (n = 158) with independent annotations across lineages, variables, and/or CNVs had functions related to mitochondrial structure/function, immune response, hypoxia, olfaction, and DNA repair. Some of these genes have been previously linked to putative high elevation and/or climate adaptation in other species, suggesting they may serve as important targets in future studies.

How Will Climate Change Affect Pikas’ Favorite Snacks?

Many animals are herbivores, which means they get all their nutrients from eating plants. American pikas are cute rabbit relatives that eat plants in the mountains. But alpine winters are harsh, so pikas spend their entire summer gathering and storing plants to eat under the winter snow. Just like people, pikas in Colorado have a favorite food: a plant called alpine avens. This plant species is a special pika snack because it contains natural preservatives called phenolics, which keep the food fresh all winter. We studied how climate change is affecting this important feature of the pika’s favorite meal. Alpine avens contains more phenolics now than it did 30 years ago, so they preserve better in storage. But there is a catch: these preservatives can be hard to digest. Studies like this help us start to understand the many complicated ways that climate change affects herbivores like pikas.

Evaluating genotyping-in-thousands by sequencing as a geneticmonitoring tool for a climate sentinel mammal using non-invasive and archival samples.

Genetic tools for wildlife monitoring can provide valuable information on spatiotemporal population trends and connectivity, particularly in systems experiencing rapid environmental change. Multiplexed targeted amplicon sequencing techniques, such as genotyping-in-thousands by sequencing (GT-seq), can provide cost-effective approaches for collecting genetic data from low-quality and quantity DNA samples, making them potentially useful for long-term wildlife monitoring using non-invasive and archival samples. Here, we developed a GT-seq panel as a potential monitoring tool for the American pika (Ochotona princeps) and evaluated its performance when applied to traditional, non-invasive, and archival samples, respectively. Specifically, we optimized a GT-seq panel (307 single nucleotide polymorphisms (SNPs)) that included neutral, sex-associated, and putatively adaptive SNPs using contemporary tissue samples (n = 77) from the Northern Rocky Mountains lineage of American pikas. The panel demonstrated high genotyping success (94.7%), low genotyping error (0.001%), and excellent performance identifying individuals, sex, relatedness, and population structure. We subsequently applied the GT-seq panel to archival tissue (n = 17) and contemporary fecal pellet samples (n = 129) collected within the Canadian Rocky Mountains to evaluate its effectiveness. Although the panel demonstrated high efficacy with archival tissue samples (90.5% genotyping success, 0.0% genotyping error), this was not the case for the fecal pellet samples (79.7% genotyping success, 28.4% genotyping error) likely due to the exceptionally low quality/quantity of recovered DNA using the approaches implemented. Overall, our study reinforced GT-seq as an effective tool using contemporary and archival tissue samples, providing future opportunities for temporal applications using historical specimens. Our results further highlight the need for additional optimization of sample and genetic data collection techniques prior to broader-scale implementation of a non-invasive genetic monitoring tool for American pikas.

Evolution of SARS-CoV-2 Spikes shapes their binding affinities to animal ACE2 orthologs.

Spike-receptor interaction is a critical determinant for the host range of coronaviruses. In this study, we investigated the SARS-CoV-2 WHU01 strain and five WHO-designated SARS-CoV-2 variants of concern (VOCs), including Alpha, Beta, Gamma, Delta, and the early Omicron variant, for their Spike interactions with ACE2 proteins of 18 animal species. First, the receptor-binding domains (RBDs) of Alpha, Beta, Gamma, and Omicron were found to display progressive gain of affinity to mouse ACE2. More interestingly, these RBDs were also found with progressive loss of affinities to multiple ACE2 orthologs. The Omicron RBD showed decreased or complete loss of affinity to eight tested animal ACE2 orthologs, including that of some livestock animals (horse, donkey, and pig), pet animals (dog and cat), and wild animals (pangolin, American pika, and Rhinolophus sinicus bat). These findings shed light on potential host range shift of SARS-CoV-2 VOCs, especially that of the Omicron variant.

Life on the edge-a changing genetic landscape within an iconic American pika metapopulation over the last half century.

Declines and extirpations of American pika (Ochotona princeps) populations at historically occupied sites started being documented in the literature during the early 2000s. Commensurate with global climate change, many of these losses at peripheral and lower elevation sites have been associated with changes in ambient air temperature and precipitation regimes. Here, we report on a decline in available genetic resources for an iconic American pika metapopulation, located at the southwestern edge of the species distribution in the Bodie Hills of eastern California, USA. Composed of highly fragmented habitat created by hard rock mining, the ore dumps at this site were likely colonized by pikas around the end of the 19th century from nearby natural talus outcrops. Genetic data extracted from both contemporary samples and archived natural history collections allowed us to track population and patch-level genetic diversity for Bodie pikas across three distinct sampling points during the last half- century (1948-1949, 1988-1991, 2013-2015). Reductions in within-population allelic diversity and expected heterozygosity were observed across the full time period. More extensive sampling of extant patches during the 1988-1991 and 2013-2015 periods revealed an increase in population structure and a reduction in effective population size. Furthermore, census records from the last 51 years as well as archived museum samples collected in 1947 from a nearby pika population in the Wassuk range (Nevada, USA) provide further support of the increasing isolation and genetic coalescence occurring in this region. This study highlights the importance of museum samples and long-term monitoring in contextualizing our understanding of population viability.

Spatio-temporal variability in the strength, directionality, and relative importance of climate on occupancy and population densities in a philopatric mammal, the American pika (Ochotona princeps)

Species distribution models (SDMs) have been widely employed to evaluate species–environment relationships. However, when extrapolated over broad spatial scales or through time, these models decline in their predictive ability due to variation in how species respond to their environment. Many models assume species–environment relationships remain constant over space and time, hindering their ability to accurately forecast distributions. Therefore, there is growing recognition that models could be improved by accounting for spatio-temporal nonstationarity – a phenomenon wherein the factors governing ecological processes change over space or time. Here, we investigated nonstationarity in American pika (Ochotona princeps) relationships with climatic variables in the Rocky Mountains (USA). We first compared broad-scale differences in pika–climate patterns for occupancy and population density across the Southern, Central, and Northern Rockies. Next, we investigated within-ecoregion variation across four mountain ranges nested within the Northern Rockies. Lastly, we tested whether species–climate relationships changed over time within the Central Rockies ecoregion. Across all analyses, we found varying levels of nonstationarity among the climate metrics for both occupancy and density. Although we found general congruence in temperature metrics, which consistently had negative coefficients, and moisture metrics (e.g., relative humidity), which had positive coefficients, nonstationarity was greatest for summer and winter precipitation over both space and time. These results suggest that interpretations from one ecoregion should not be applied to other regions universally – especially when using precipitation metrics. The within-ecoregion analysis found much greater variation in the strength-of-relationship coefficients among the four mountain ranges, relative to the inter-regional analysis, possibly attributable to smaller sample sizes per mountain range. Lastly, the importance of several variables shifted through time from significant to insignificant in the temporal analysis. Our results collectively reveal the overall complexity underlying species–environment relationships. With rapidly shifting conditions globally, this work adds to the growing body of literature highlighting how issues of spatio-temporal nonstationarity can limit the accuracy, transferability, and reliability of models and that interpretations will likely be most robust at local to regional scales. Diagnosing, describing, and incorporating nonstationarity of species–climate relationships into models over space and time could serve as a pivotal step in creating more informative models.

Integrating multiple sign types to improve occupancy estimation for inconspicuous species.

Standard occupancy models enable unbiased estimation of occupancy by accounting for observation errors such as missed detections (false negatives) and, less commonly, incorrect detections (false positives). Occupancy models are fitted to data from repeated site visits in which surveyors record evidence of species presence. Use of indirect sign (e.g., scat, tracks) as evidence of presence can vastly improve survey efficiency for inconspicuous species but can also introduce additional sources of error. We developed a "multi-sign" occupancy approach to model the detection process separately for unique sign types and used this method to improve estimates of occupancy dynamics for an inconspicuous species, the American pika (Ochotona princeps). We investigated how estimates of pika occupancy and environmental drivers differed under four increasingly realistic representations of the observation process: (1) perfect detection (commonly assumed for modeling pika occupancy), (2) standard occupancy model (single observation process without possibility of false detection), (3) multi-sign with no false detections (non-false positive model), and (4) multi-sign with false detections (full model). For the multi-sign occupancy models, we modeled the detection of each sign type (fresh scat, fresh haypiles, pika calls, and pika sightings) separately as a function of climatic and environmental covariates. Estimates of occupancy processes and inferences about environmental drivers were sensitive to different detection models. Simplified representations of the detection processes generally resulted in higher occupancy estimates and higher turnover rates than the full multi-sign model. Environmental drivers also varied in their influence on occupancy models, where (e.g.) forb cover was estimated to more strongly influence occupancy in the full multi-sign model than the simpler models. As has been reported previously in other contexts, unmodeled heterogeneity in the observation process can lead to biases in occupancy processes and uncertainty in the relationships between occupancy and environmental covariates. Overall, our multi-sign approach to dynamic occupancy modeling, which accounts for spatio-temporal variation in reliability among sign types, has strong potential to generate more realistic estimates of occupancy dynamics for inconspicuous species.

Sampling a pika's pantry: Temporal shifts in nutritional quality and winter preservation of American pika food caches

AbstractClimate change is increasing temperature, decreasing precipitation, and increasing atmospheric CO2 concentrations in many ecosystems. As atmospheric carbon rises, plants may increase carbon‐based defenses, such as phenolics, thereby potentially affecting food quality, foraging habits, and habitat suitability for mammalian herbivores. In alpine habitats, the American pika (Ochotona princeps) is a model species for studying effects of changing plant chemistry on mammals. To survive between growing seasons, pikas cache “haypiles” of plants rich in phenolics. Although they are toxic to pikas, phenolic compounds facilitate retention of plant biomass and nutrition during storage, and they degrade over time. Alpine avens (Geum rossii, Rosales: Rosaceae) is a high‐phenolic plant species that comprises up to 75% of pika haypiles in Colorado. Here, we tested the hypothesis that contemporary climate change has affected the nutritional value of alpine avens to pikas in the last 30 years. Specifically, we compared phenolic activity, nutritional quality, and overwinter preservation of plants collected at Niwot Ridge, Colorado (USA), in 1992 to those collected between 2010 and 2018, spanning nearly three decades of climate change. Phenolic activity increased in alpine avens since 1992, while fiber and nitrogen content decreased. Importantly, overwinter preservation of plant biomass also increased, particularly on windblown slopes without long‐lasting snow cover. Previous studies indicate that pikas at this site still depend on alpine avens for their winter food caches. Higher phenolic content in alpine avens could therefore enhance the preservation of haypiles over winter; however, if pikas must further delay consuming these plants to avoid toxicity or invest extra energy in detoxification, then the nutritional gains from enhanced preservation may not be beneficial. This study provides important insights into how climate‐driven changes in plant chemistry will affect mammalian herbivores in the future.

Geographic and taxonomic variation in adaptive capacity among mountain-dwelling small mammals: Implications for conservation status and actions

Contemporary climate change is modifying the distribution, morphology, phenology, physiology, evolution, and interspecific interactions of species. Effects of climate change are mediated not only through the magnitude of change experienced (exposure) and an animal's sensitivity to such changes, but also through the ability of the population or species to adjust to climatic variability and change genetically, behaviorally, or spatially (via its distribution) (i.e., adaptive capacity; AC). Here, we used an attribute-based framework to systematically evaluate and compare the AC of American pikas (Ochotona princeps) against four other mountain-dwelling small mammals of North America to determine whether pikas are disproportionately vulnerable to climate change, as has been postulated. Unlike previous analyses, we also compared AC across O. princeps lineages and across three taxonomic (and thus, spatial) scales. Our results indicate that pikas have markedly lower adaptive capacity than all compared species except bushy-tailed woodrats (Neotoma cinerea), and that our assessments of species generally align with earlier characterizations of climate-change vulnerability based on life-history characteristics. Although AC did not differ dramatically among pika lineages, some attributes are likely constraining AC differently in various parts of the geographic range. Comparisons across taxonomic levels of pikas illustrated that, although AC levels were comparable in pika lineages versus range-wide, AC was assessed as lower in interior-Great-Basin pikas than across the entire O.p. schisticeps lineage. We conclude that the comparatively lower AC of pikas results in particularly high susceptibility to anthropogenic climate change, corroborating results from numerous other recent investigations of pikas' climate-responsiveness. Adaptive-capacity evaluations appear useful as a consistent way to identify sentinel species or populations and for conservation prioritization.

Stress-associated metabolites vary with both season and habitat across populations of a climate sentinel species

ABSTRACT Relating physiological stress to habitat quality could refine conservation efforts. Habitat quality, which is often inferred from patch occupancy or demographic rates, might be measured in a more timely and nuanced way using metrics of physiological stress. To understand whether stress-associated hormones vary with metrics of habitat quality, we measured fecal glucocorticoid metabolite (FGM) levels in the American pika (Ochotona princeps), a small mammal with well-defined habitat (talus), which can vary in quality depending on the presence of subsurface ice features. In spring and fall 2018, we collected feces noninvasively from pika territories in taluses “with” or “without” subsurface ice to capture seasonal variation in FGM between habitat types. We used linear mixed effects models to explore the interactions among season, habitat metrics (including subsurface ice status), and subsurface temperature as predictors of FGM. We found support for interacting effects on FGM levels, which covaried with season, elevation, putative ice presence, graminoid to forb ratio, graminoid cover, and measures of acute subsurface heat exposure. However, only one subsurface temperature metric differed according to putative presence of subsurface ice. Our results contribute to the growing evidence that FGMs might be developed as a tool to assess habitat quality.

Anatomical Correlates of Cursoriality are Compromised by Body Size and Propensity to Burrow in a Group of Small Mammals (Lagomorpha)

Highly cursorial animals are specialised for fast, sustained running via specific morphological adaptations, notably including changes in limb segment length and mechanical advantage. Members of the order Lagomorpha (hares, rabbits and pikas) vary in cursorial ability; hares are generally highly cursorial, rabbits more frequently saltate, and pikas predominantly trot. Previous investigations of lagomorphs have identified anatomical trends correlated with this ‘cursoriality gradient’, however, the phylogenetic sampling of such investigations has been limited to three American species, namely the American pika (Ochotona princeps), brush rabbit (Sylvilagus bachmani), and black-tailed jackrabbit (Lepus californicus). Here, we expand the phylogenetic sample and body size range by including novel data from Australian samples of the European rabbit (Oryctolagus cuniculus) and European hare (L. europaeus), alongside unpublished data on the Eastern cottontail (S. floridanus). X-ray Computed Tomography and digital landmarking were used to capture proportions within the appendicular skeleton of ~ 40 specimens of each European species. In doubling the number of species studied, we find the previously-identified morphological gradients associated with cursorial behaviour are complicated when evaluated in the larger sample. The relative length and joint velocity of limbs was found to be lower than predicted in European rabbits and hares. Furthermore, we present a novel assessment of morphological integration in the lagomorph appendicular skeleton, finding between-limb covariation patterns that are generally similar to those of other mammals. Broadly, these results suggest cursoriality is only one of many selective forces driving lagomorph skeletal evolution, with variations in body size and fossoriality potentially having measurable impacts.

Return of the pika: American pikas re-occupy long-extirpated, warm locations.

American pikas (Ochotona princeps), small mammals related to rabbits, occur in mountainous regions of western North America, where they live in shattered‐rock habitats (talus). Aspects of their physiology and life history create situations that appear to put pikas at risk from warming climates. Some low‐elevation, warm sites that historically harbored pikas have become extirpated, and the assumption is that these will not be re‐colonized under current climate trends. Unexpectedly, in 2021, we found that pikas had re‐colonized two very warm, low‐elevation, dry sites in eastern California, USA, in the Bodie Mountains and Mono Craters. Resident pikas appear to have been absent at both sites for ≥10 years. These findings suggest that pikas, which are normally diurnally active, are able to overcome thermal dispersal barriers and re‐colonize long‐extirpated sites, perhaps by moving during cool nights. Our data also highlight the often unrecognized suitability of pika habitat in warm regions where the interiors of taluses can remain stably cool even when external air temperatures are hot.

Revisiting talus and free-air temperatures after 50 years of change at an American pika (Ochotona princeps) study site in the Southern Rockies

Climate change in mountain regions has exposed high-elevation species to rapidly changing temperatures. Although climate exposure can be reduced in certain microclimates, the quality of microclimatic refugia might also degrade with climate change. The American pika (Ochotona princeps) often inhabits high elevations, and is considered climate-sensitive due to its narrow thermal tolerance and recent extirpations in some warmer portions of its range. Pikas behaviorally thermoregulate by taking refuge in the subsurface microclimates found in taluses and other rocky habitats, where daily thermal fluctuations are attenuated and somewhat decoupled from free-air temperatures. Changes in microclimate might reduce the efficacy of this behavioral thermoregulation. This study compares recent (2009–2021) subsurface temperatures at a long-term pika study site with a rare instance of historical (1963–1964) data from the same location. We also place historical and recent microclimates in context using long-term data on free-air temperatures from the same area. Recent free-air temperatures were often warmer than historical records, and subsurface temperatures exhibited even stronger warming between periods. Temperatures measured in the talus were often dramatically warmer in recent records, especially at the deeper of two subsurface sensor placements in this study. Winter months showed the greatest changes in both talus and free-air temperatures. Differences between historical and recent microclimates were not explained by the precise placement of sensors, as recent temperatures were similar across a wide variety of subsurface placements, and temporal changes in free-air temperatures at the historical study site were also reflected in data from nearby weather stations. Together, these results suggest that subsurface microclimates important for pika thermoregulation have changed over the past few decades, perhaps even faster than observed changes in free-air temperatures. The generality of these results and their potential ramifications for ecosystem processes and services should be explored.

From Habitat to Hormones: Year-around territorial behavior in rock-dwelling but not in forest and grassland lagomorphs and the role of DHEA

Year-around defense of extremely patchy habitat may require hormones that drive territorial behavior, but have no other costly physiological effects. The American pika (Ochotona princeps), but not the snowshoe hare (Lepus americanus) nor the eastern cottontail rabbit (Sylvilagus floridanus), exhibits this behavior. The former engages in contest competition against all individuals independent of sex to protect its territory in highly fragmented patches on mountain talus slopes; the latter in scramble competition in more continuous forests, grasslands, and shrublands. The hormonal basis for this difference in lagomorphs is unknown. Dehydroepiandrosterone is a prohormone produced by the zona reticularis of the adrenal cortex. It has no effect on aggressive behavior until converted in the brain to estrogen. We assessed levels (DHEA-S plus DHEA) in all species collected in the wild. In nonbreeding pikas, levels were 256 times higher than in hares and 22 times higher than in rabbits. Within species, females and males had similar levels. The proportion of the adrenal cortex devoted to the zona reticularis was significantly larger in pikas than in hares or rabbits. Our evidence is consistent with the hypothesis that dehydroepiandrosterone drives this individual-based, year-around territoriality in pikas. We propose a definitive experiment to determine this and recommend comparative studies in central Asia where there is high diversity of pika species whose behavior ranges from individual-based territoriality to colonial. Thus, we speculate that the wild American pika has the adrenal-brain nexus for all seasons and is an excellent model to understand how habitat drives the hormonal control of spacing behavior.

Life-history traits and fitness of plateau pika (Ochotona curzoniae) in alpine meadow ecosystem.

Plateau pika (Ochotona curzoniae) is an endemic mammal living in the alpine meadow ecosystem in Qinghai–Tibet Plateau. We studied life history of plateau pika by mark–recapturing method. The research results showed that fitness of plateau pika to its habitat was lower than those of many other mammal species; plateau pika adopted quick growth strategy; the life‐history features of plateau pika were accorded with Charnov (Evolutionary Ecology Research, 2002, 4, 749)'s formula about life‐history classification, that is, E/α ≈ 1.35, C·E ≈ 1.7, I/m ≈ 0.3, in which E stands for average adult life span; α stands for age at first reproduction; C is reproductive effort; I is the size of an offspring at independence from the parent; m is average adult body mass. There does not exist dimorphism in plateau pikas. The configurations of females and males are similar; the average body mass of males is a little heavier than that of females, but the difference is not significant (F = 1.0854, df = 154, p > .3058). The juveniles exhibit a J‐form growth curve before 30‐day age and grow slower between 30 and 65‐day age and reach body mass equilibrium at about 65 days old. So, 65 days is the maturity age of plateau pika. There are 3 mortality peaks in plateau pika population. The first time is in neonate period, when only one half of juveniles can pass through this period, which implies that juvenile period is influenced strongly by natural selection; the second time is in fecundity peak period, which reflects the cost of reproduction; the third time is in the old age of plateau pika, a significant loss occurs during this period, which is the result of natural selection. The average longevity of females is longer than that of males. A female reaches the maximum life span recorded, that is, 931 days. The average longevity of all individuals is 16.33 months ≈ 490 days. The survival rate of females is higher than that of males, which reflects the cost of reproduction and society role of males. It is the outcome of natural selection. The sex ratio of neonates is 1:1; however, the sex ratio of adults is female: male = 1.31:1, which is caused by higher mortality of males over females in life history. Plateau pikas reproduced two times every year. The average gestation period of females is 18–20 days. The average litter size is 4.57 individual. The average body mass of neonates is 9.28 g. The average litter size of adult female plateau pika does not variate with age. Breeding season is between April and June. The reproductive value and fertility of 15–18 months age females are highest. The reproductive value and fertility increased with age before reproductive value and fertility peak age; however, the reproductive value and fertility decreased with age after peak age. The fitness of plateau pika (r = .1125) was lower than that of American pikas (O. princeps) (r = 2.172). The survival rate was the main factor influencing fitness. The dynamic trend of plateau pika population was coincident with r, that is, the plateau pika population was stable.

On pikas and ticks

The pika (Ochotona princeps) is a lagomorph that typically lives in rocky areas such as talus slopes, at elevations of 8000 to 13,000 feet in the Rocky Mountains of western North America. As the climate warms, species from lower elevations can shift their range upwards, but most pika populations have nowhere to go because they already live at the highest elevations (eg J Mammal 2020; doi.org/10.1093/jmammal/gyaa110). Range shifts of species from lower elevations might also harm pikas. In October 2020, one of us (Olmstead) photographed this pika near Silver Lake in Big Cottonwood Canyon, in the Wasatch Range within Salt Lake County, Utah. Numerous engorged ticks (species unknown) are evident in the lesion on its throat, and others appear to be around its ear. Johanna Varner, a pika expert at Colorado Mesa University (Grand Junction, CO), remarked: “None of us [referring to herself and other authorities] have ever seen a pika with so many ticks on it: in fact, your pika had more ticks on its cheek and ear than I’ve seen on all pikas combined in my 10 years of watching them!” Various tick species have expanded their ranges as the climate has warmed. If these and other ectoparasites become more prevalent in alpine zones, then the health of alpine wildlife populations, such as those of the already vulnerable pika, could be adversely affected.