Pursuit and escape drive fine-scale movement variation during migration in a temperate alpine ungulate.

Climate change reduces snowpack, advances snowmelt phenology, drives summer warming, alters growing season precipitation regimes, and consequently modifies vegetation phenology in mountain systems. Elevational migrants track spatial variation in seasonal plant growth by moving between ranges at different elevations during spring, so climate-driven vegetation change may disrupt historic benefits of migration. Elevational migrants can furthermore cope with short-term environmental variability by undertaking brief vertical movements to refugia when sudden adverse conditions arise. We uncover drivers of fine-scale vertical movement variation during upland migration in an endangered alpine specialist, Sierra Nevada bighorn sheep (Ovis canadensis sierrae) using a 20-year study of GPS collar data collected from 311 unique individuals. We used integrated step-selection analysis to determine factors that promote vertical movements and drive selection of destinations following vertical movements. Our results reveal that relatively high temperatures consistently drive uphill movements, while precipitation likely drives downhill movements. Furthermore, bighorn select destinations at their peak annual biomass and maximal time since snowmelt. These results indicate that although Sierra Nevada bighorn sheep seek out foraging opportunities related to landscape phenology, they compensate for short-term environmental stressors by undertaking brief up- and downslope vertical movements. Migrants may therefore be impacted by future warming and increased storm frequency or intensity, with shifts in annual migration timing, and fine-scale vertical movement responses to environmental variability.

Infection-nutrition feedbacks: fat supports pathogen clearance but pathogens reduce fat in a wild mammal.

Though far less obvious than direct effects (clinical disease or mortality), the indirect influences of pathogens are difficult to estimate but may hold fitness consequences. Here, we disentangle the directional relationships between infection and energetic reserves, evaluating the hypotheses that energetic reserves influence infection status of the host and that infection elicits costs to energetic reserves. Using repeated measures of fat reserves and infection status in individual bighorn sheep (Ovis canadensis) in the Greater Yellowstone Ecosystem, we documented that fat influenced ability to clear pathogens (Mycoplasma ovipneumoniae) and infection with respiratory pathogens was costly to fat reserves. Costs of infection approached, and in some instances exceeded, costs of rearing offspring to independence in terms of reductions to fat reserves. Fat influenced probability of clearing pathogens, pregnancy and over-winter survival; from an energetic perspective, an animal could survive for up to 23 days on the amount of fat that was lost to high levels of infection. Cost of pathogens may amplify trade-offs between reproduction and survival. In the absence of an active outbreak, the influence of resident pathogens often is overlooked. Nevertheless, the energetic burden of pathogens likely has consequences for fitness and population dynamics, especially when food resources are insufficient.

The abundance and persistence of Caprinae populations.

Stable or growing populations may go extinct when their sizes cannot withstand large swings in temporal variation and stochastic forces. Hence, the minimum abundance threshold defining when populations can persist without human intervention forms a key conservation parameter. We identify this threshold for many populations of Caprinae, typically threatened species lacking demographic data. Doing so helps triage conservation and management actions for threatened or harvested populations. Methodologically, we used population projection matrices and simulations, with starting abundance, recruitment, and adult female survival predicting future abundance, growth rate (λ), and population trend. We incorporated mean demographic rates representative of Caprinae populations and corresponding variances from desert bighorn sheep (Ovis canadensis nelsoni), as a proxy for Caprinae sharing similar life histories. We found a population's minimum abundance resulting in ≤ 0.01 chance of quasi-extinction (QE; population ≤ 5 adult females) in 10 years and ≤ 0.10 QE in 30 years as 50 adult females, or 70 were translocation (removals) pursued. Discovering the threshold required 3 demographic parameters. We show, however, that monitoring populations' relationships to this threshold requires only abundance and recruitment data. This applied approach avoids the logistical and cost hurdles in measuring female survival, making assays of population persistence more practical.

1H NMR based metabolic profiling distinguishes the differential impact of capture techniques on wild bighorn sheep.

Environmental metabolomics has the potential to facilitate the establishment of a new suite of tools for assessing the physiological status of important wildlife species. A first step in developing such tools is to evaluate the impacts of various capture techniques on metabolic profiles as capture is necessary to obtain the biological samples required for assays. This study employed 1H nuclear magnetic resonance (NMR)-based metabolite profiling of 562 blood serum samples from wild bighorn sheep to identify characteristic molecular serum makers of three capture techniques (dart, dropnet, and helicopter-based captures) to inform future sampling protocols for metabolomics studies, and to provide insights into the physiological impacts of capture. We found that different capture techniques induce distinct changes in amino acid serum profiles, the urea cycle, and glycolysis, and attribute the differences in metabolic patterns to differences in physical activity and stress caused by the different capture methods. These results suggest that when designing experiments involving the capture of wild animals, it may be prudent to employ a single capture technique to reduce confounding factors. Our results also supports administration of tranquilizers as soon as animals are restrained to mitigate short-term physiological and metabolic responses when using pursuit and physical restraint capture techniques.

Bioinspired material architectures from bighorn sheep horncore velar bone for impact loading applications.

Rocky Mountain bighorn sheep rams (Ovis canadensis canadensis) routinely conduct intraspecific combat where high energy cranial impacts are experienced. Previous studies have estimated cranial impact forces to be up to 3400 N during ramming, and prior finite element modeling studies showed the bony horncore stores 3 × more strain energy than the horn during impact. In the current study, the architecture of the porous bone within the horncore was quantified, mimicked, analyzed by finite element modeling, fabricated via additive manufacturing, and mechanically tested to determine the suitability of the novel bioinspired material architecture for use in running shoe midsoles. The iterative biomimicking design approach was able to tailor the mechanical behavior of the porous bone mimics. The approach produced 3D printed mimics that performed similarly to ethylene-vinyl acetate shoe materials in quasi-static loading. Furthermore, a quadratic relationship was discovered between impact force and stiffness in the porous bone mimics, which indicates a range of stiffness values that prevents impact force from becoming excessively high. These findings have implications for the design of novel bioinspired material architectures for minimizing impact force.

Year-round water management for desert bighorn sheep corresponds with visits by predators not bighorn sheep.

Managing water (e.g., catchments) to increase the abundance and distribution of game is popular in arid regions, especially throughout the southwest United States, where biologists often manage water year-round for desert bighorn sheep (Ovis canadensis nelsoni). Bighorn may visit water when predators (e.g., mountain lions [Puma concolor], coyotes [Canis latrans]) do not, suggesting that differences in species ecology or their surface water requirements influence visit timing. Alternatively, visits by desert bighorn sheep and predators may align. The former outcome identifies opportunities to improve water management by providing water when desert bighorn sheep visit most, which hypothetically may reduce predator presence, range expansion and predation, thereby supporting objectives to increase sheep abundances. Since advancing water management hinges on understanding the patterns of species visits, we identified when these three species and mule deer (Odocoileus hemionus) visited managed waters in three North American deserts (Chihuahuan, Sonoran, Mojave). We unraveled the ecological basis describing why visits occurred by associating species visits with four weather variables using multi-site, multi-species models within a Bayesian hierarchical framework (3.4 million images; 105 locations; 7/2009-12/2016). Desert bighorn sheep concentrated visits to water within 4-5 contiguous months. Mountain lions visited water essentially year-round within all deserts. Higher maximum temperature influenced visits to water, especially for desert bighorn sheep. Less long-term precipitation (prior 6-week total) raised visits for all species, and influenced mountain lion visits 3-20 times more than mule deer and 3-37 times more than sheep visits. Visits to water by prey were inconsistent predictors of visits to water by mountain lions. Our results suggest improvements to water management by aligning water provision with the patterns and ecological explanations of desert bighorn sheep visits. We exemplify a scientific approach to water management for enhancing stewardship of desert mammals, be it the southwest United States or arid regions elsewhere.

Sons accelerate maternal aging in a wild mammal.

Aging, or senescence, is a progressive deterioration of physiological function with age. It leads to age-related declines in reproduction (reproductive senescence) and survival (actuarial senescence) in most organisms. However, senescence patterns can be highly variable across species, populations, and individuals, and the reasons for such variations remain poorly understood. Evolutionary theories predict that increases in reproductive effort in early life should be associated with accelerated senescence, but empirical tests have yielded mixed results. Although in sexually size-dimorphic species offspring of the larger sex (typically males) commonly require more parental resources, these sex differences are not currently incorporated into evolutionary theories of aging. Here, we show that female reproductive senescence varies with both the number and sex ratio of offspring weaned during early life, using data from a long-term study of bighorn sheep. For a given number of offspring, females that weaned more sons than daughters when aged between 2 and 7 y experienced faster senescence in offspring survival in old age. By contrast, analyses of actuarial senescence showed no cost of early-life reproduction. Our results unite two important topics in evolutionary biology: life history and sex allocation. Offspring sex ratio may help explain among-individual variation in senescence rates in other species, including humans.

Habitat use and social mixing between groups of resident and augmented bighorn sheep.

Monitoring dispersal, habitat use, and social mixing of released ungulates is crucial for successful translocation and species conservation. We monitored 127 female bighorn sheep (Ovis canadensis) released in three populations from 2000 to 2009 to investigate if augmented bighorns expanded and shifted seasonal ranges, used different habitat compared with resident females, and if animals mixed socially. Augmented bighorns in all populations expanded range use compared with residents by shifting utilization distributions. Size of utilization distributions, however, were smaller for augmented females compared with residents in all areas except one. Overlap of seasonal utilization distributions between augmented and resident bighorns and use of slope and elevation differed across populations. In two populations, differences in size and overlap of seasonal utilization distributions and use of slope and elevation supported the hypothesis that habitat use of bighorns in their source area influenced their habitat use after release. Mixing between resident and augmented adult females occurred on average during only 21% of sightings and was similar across populations. Our results clarify how augmented bighorns mix with resident animals and how habitat use is modified following augmentations. Such information is needed to improve bighorn sheep augmentations and can be applied to augmentations of other ungulates.

Maternal longevity and offspring sex in wild ungulates.

In species with sexual size dimorphism, the offspring of the larger sex usually have greater energy requirements and may lead to greater fitness costs for parents. The effects of offspring sex on maternal longevity, however, have only been tested in humans. Human studies produced mixed results and considerable debate mainly owing to the difficulty of distinguishing the effects of sexual dimorphism from sociocultural factors. To advance this debate, we examined how the relative number of sons influenced maternal longevity in four species of free-living ungulates (Soay sheep Ovis aries; bighorn sheep, Ovis canadensis; red deer, Cervus elaphus; mountain goat, Oreamnos americanus), with high male-biased sexual size dimorphism but without complicating sociocultural variables. We found no evidence for a higher cumulative cost of sons than of daughters on maternal longevity. For a given number of offspring, most females with many sons in all four populations lived longer than females with few sons. The higher cost of sons over daughters on maternal lifespan reported by some human studies may be the exception rather than the rule in long-lived iteroparous species.

Waterhole detection using a vegetation index in desert bighorn sheep (Ovis canadensis cremnobates) habitat.

In arid ecosystems, desert bighorn sheep are dependent on natural waterholes, particularly in summer when forage is scarce and environmental temperatures are high. To detect waterholes in Sierra Santa Isabel, which is the largest area of desert bighorn sheep habitat in the state of Baja California, Mexico, we used the normalized difference vegetation index (NDVI) and normalized difference water index (NDWI) from Sentinel-2 satellite images. Waterhole detection was based on the premise that sites with greater water availability, where NDVI was higher, can be identified by their density of vegetation greenness. For the detected waterholes, we estimated the escape terrain (presence of cliffs or steep, rocky slopes) around each by the vector ruggedness measure to determine their potential use by desert bighorn sheep based on the animals' presence as documented by camera traps. We detected 14 waterholes with the NDVI of which 11 were known by land owners and 3 were unrecorded. Desert bighorn were not detected in waterholes with high values of escape terrain, i.e., flat areas. Waterhole detection by NDVI is a simple method, and with the assistance and knowledge of the inhabitants of the Sierra, it was possible to confirm the presence each waterhole in the field.

Effects of intrinsic and extrinsic factors on ruminating, grazing, and bedding time in bighorn sheep (Ovis canadensis).

Rumination is the repeated process of regurgitation of a food bolus, followed by chewing, swallowing, and regurgitation, which enhance nutrient assimilation. Time spent in lateral recumbency (i.e., bedded, lying) has often been used as a proxy for time spent ruminating due to difficulties of observing detailed rumination behavior in the field. The actual proportion of time spent ruminating, or other activities, will in turn be affected by the age and sex of an individual but also with changes in food quality. We studied the effects of intrinsic and extrinsic factors on time spent ruminating, bedding, proportion of bedding time spent ruminating, and grazing of individually marked bighorn sheep (Ovis canadensis). Our results show that bighorn sheep spent more time ruminating and less time grazing in summer and autumn. Overall, females spent less time ruminating, and more time grazing than males. Bighorn sheep decreased their time spent ruminating with increasing acid detergent fiber (ADF) content in the forage. Age influenced the time spent grazing, bedded and proportion of bedded time spent ruminating. Older sheep not only increased their bedding time but also their time spent bedded without ruminating compared to younger individuals. The proportion of time spent grazing was also affected by age, with a decrease in the proportion of time spent grazing with increasing age. Our results suggest that these four behaviors are plastic and variable. We thus conclude that bedding time does not reflect time spent ruminating but that the latter is affected by both intrinsic and extrinsic factors.

Niche similarities among introduced and native mountain ungulates.

The niche concept provides a strong foundation for theoretical and applied research among a broad range of disciplines. When two ecologically similar species are sympatric, theory predicts they will occupy distinct ecological niches to reduce competition. Capitalizing on the increasing availability of spatial data, we built from single species habitat suitability models to a multispecies evaluation of the niche partitioning hypothesis with sympatric mountain ungulates: native bighorn sheep (BHS; Ovis canadensis) and introduced mountain goats (MTG; Oreamnos americanus) in the northeast Greater Yellowstone Area. We characterized seasonal niches using two-stage resource selection functions with a used-available design and descriptive summaries of the niche attributes associated with used GPS locations. We evaluated seasonal similarity in niche space according to confidence interval overlap of model coefficients and similarity in geographic space by comparing model predicted values with Schoener's D metric. Our sample contained 37,962 summer locations from 53 individuals (BHS = 31, MTG = 22), and 79,984 winter locations from 57 individuals (BHS = 35, MTG = 22). Slope was the most influential niche component for both species and seasons, and showed the strongest evidence of niche partitioning. Bighorn sheep occurred on steeper slopes than mountain goats in summer and mountain goats occurred on steeper slopes in winter. The pattern of differential selection among species was less prevalent for the remaining covariates, indicating similarity in niche space. Model predictions in geographic space showed broad seasonal similarity (summer D = 0.88, winter D = 0.87), as did niche characterizations from used GPS locations. The striking similarities in seasonal niches suggest that introduced mountain goats will continue to increase their spatial overlap with native bighorn. Our results suggest that reducing densities of mountain goats in hunted areas where they are sympatric with bighorn sheep and impeding their expansion may reduce the possibility of competition and disease transfer. Additional studies that specifically investigate partitioning at finer scales and along dietary or temporal niche axes will help to inform an adaptive management approach.

Direct and indirect effects of early-life environment on lifetime fitness of bighorn ewes.

Cohort effects, when a common environment affects long-term performance, can have a major impact on population dynamics. Very few studies of wild animals have obtained the necessary data to study the mechanisms leading to cohort effects. We exploited 42 years of individual-based data on bighorn sheep to test for causal links between birth density, body mass, age at first reproduction (AFR), longevity and lifetime reproductive success (LRS) using path analysis. Specifically, we investigated whether the effect of early-life environment on lifetime fitness was the result of indirect effects through body mass or direct effects of early-life environment on fitness. Additionally, we evaluated whether the effects of early-life environment were dependant on the environment experienced during adulthood. Contrary to expectation, the effect on LRS mediated through body mass was weak compared to the effects found via a delay in AFR, reduced longevity and the direct effect of birth density. Birth density also had an important indirect effect on LRS through reduced longevity, but only when adult density was high. Our results show that the potential long-term consequences of a harsh early-life environment on fitness are likely to be underestimated if investigations are limited to body mass instead of fitness at several life stages, or if the interactions between past and present environment are ignored.

Long-term fitness consequences of early environment in a long-lived ungulate.

Cohort effects can be a major source of heterogeneity and play an important role in population dynamics. Silver-spoon effects, when environmental quality at birth improves future performance regardless of the adult environment, can induce strong lagged responses on population growth. Alternatively, the external predictive adaptive response (PAR) hypothesis predicts that organisms will adjust their developmental trajectory and physiology during early life in anticipation of expected adult conditions but has rarely been assessed in wild species. We used over 40 years of detailed individual monitoring of bighorn ewes (Ovis canadensis) to quantify long-term cohort effects on survival and reproduction. We then tested both the silver-spoon and the PAR hypotheses. Cohort effects involved a strong interaction between birth and current environments: reproduction and survival were lowest for ewes that were born and lived at high population densities. This interaction, however, does not support the PAR hypothesis because individuals with matching high-density birth and adult environments had reduced fitness. Instead, individuals born at high density had overall lower lifetime fitness suggesting a silver-spoon effect. Early-life conditions can induce long-term changes in fitness components, and their effects on cohort fitness vary according to adult environment.

Paternal reproductive success drives sex allocation in a wild mammal.

Parents should bias sex allocation toward offspring of the sex most likely to provide higher fitness returns. Trivers and Willard proposed that for polygynous mammals, females should adjust sex-ratio at conception or bias allocation of resources toward the most profitable sex, according to their own body condition. However, the possibility that mammalian fathers may influence sex allocation has seldom been considered. Here, we show that the probability of having a son increased from 0.31 to 0.60 with sire reproductive success in wild bighorn sheep (Ovis canadensis). Furthermore, our results suggest that females fertilized by relatively unsuccessful sires allocated more energy during lactation to daughters than to sons, while the opposite occurred for females fertilized by successful sires. The pattern of sex-biased offspring production appears adaptive because paternal reproductive success reduced the fitness of daughters and increased the average annual weaning success of sons, independently of maternal allocation to the offspring. Our results illustrate that sex allocation can be driven by paternal phenotype, with profound influences on the strength of sexual selection and on conflicts of interest between parents.

Sex-based differences in the adaptive value of social behavior contrasted against morphology and environment.

The adaptive nature of sociality has long been a central question in ecology and evolution. However, the relative importance of social behavior for fitness, compared to morphology and environment, remains largely unknown. We assessed the importance of sociality for fitness (lamb production and survival) in a population of mark6d bighorn sheep (Ovis canadensis) over 16 years (n = 1022 sheep-years). We constructed social networks from observations (n = 38,350) of group membership (n = 3150 groups). We then tested whether consistent individual differences in social behavior (centrality) exist and evaluated their relative importance compared to factors known to affect fitness: mass, age, parental effects, and population density. Sheep exhibited consistent individual differences in social centrality. Controlling for maternal carryover effects and age, the positive effect of centrality in a social network on adult female lamb production and survival was equal or greater than the effect of body mass or population density. Social centrality had less effect on male survival and no effect on adult male lamb production or lamb survival. Through its effect on lamb production and survival, sociality in fission-fusion animal societies may ultimately influence population dynamics equally or more than morphological or environmental effects.

Comparing measures of breeding inequality and opportunity for selection with sexual selection on a quantitative character in bighorn rams.

The reliability and consistency of the many measures proposed to quantify sexual selection have been questioned for decades. Realized selection on quantitative characters measured by the selection differential i was approximated by metrics based on variance in breeding success, using either the opportunity for sexual selection Is or indices of inequality. There is no consensus about which metric best approximates realized selection on sexual characters. Recently, the opportunity for selection on character mean OSM was proposed to quantify the maximum potential selection on characters. Using 21 years of data on bighorn sheep (Ovis canadensis), we investigated the correlations between seven indices of inequality, Is , OSM and i on horn length of males. Bighorn sheep are ideal for this comparison because they are highly polygynous and sexually dimorphic, ram horn length is under strong sexual selection, and we have detailed knowledge of individual breeding success. Different metrics provided conflicting information, potentially leading to spurious conclusions about selection patterns. Iδ, an index of breeding inequality, and, to a lesser extent, Is showed the highest correlation with i on horn length, suggesting that these indices document breeding inequality in a selection context. OSM on horn length was strongly correlated with i, Is and indices of inequality. By integrating information on both realized sexual selection and breeding inequality, OSM appeared to be the best proxy of sexual selection and may be best suited to explore its ecological bases.

Costs and benefits of group living with disease: a case study of pneumonia in bighorn lambs (Ovis canadensis).

Group living facilitates pathogen transmission among social hosts, yet temporally stable host social organizations can actually limit transmission of some pathogens. When there are few between-subpopulation contacts for the duration of a disease event, transmission becomes localized to subpopulations. The number of per capita infectious contacts approaches the subpopulation size as pathogen infectiousness increases. Here, we illustrate that this is the case during epidemics of highly infectious pneumonia in bighorn lambs (Ovis canadensis). We classified individually marked bighorn ewes into disjoint seasonal subpopulations, and decomposed the variance in lamb survival to weaning into components associated with individual ewes, subpopulations, populations and years. During epidemics, lamb survival varied substantially more between ewe-subpopulations than across populations or years, suggesting localized pathogen transmission. This pattern of lamb survival was not observed during years when disease was absent. Additionally, group sizes in ewe-subpopulations were independent of population size, but the number of ewe-subpopulations increased with population size. Consequently, although one might reasonably assume that force of infection for this highly communicable disease scales with population size, in fact, host social behaviour modulates transmission such that disease is frequency-dependent within populations, and some groups remain protected during epidemic events.

A spatial risk assessment of bighorn sheep extirpation by grazing domestic sheep on public lands.

Bighorn sheep currently occupy just 30% of their historic distribution, and persist in populations less than 5% as abundant overall as their early 19th century counterparts. Present-day recovery of bighorn sheep populations is in large part limited by periodic outbreaks of respiratory disease, which can be transmitted to bighorn sheep via contact with domestic sheep grazing in their vicinity. In order to assess the viability of bighorn sheep populations on the Payette National Forest (PNF) under several alternative proposals for domestic sheep grazing, we developed a series of interlinked models. Using telemetry and habitat data, we characterized herd home ranges and foray movements of bighorn sheep from their home ranges. Combining foray model movement estimates with known domestic sheep grazing areas (allotments), a Risk of Contact Model estimated bighorn sheep contact rates with domestic sheep allotments. Finally, we used demographic and epidemiologic data to construct population and disease transmission models (Disease Model), which we used to estimate bighorn sheep persistence under each alternative grazing scenario. Depending on the probability of disease transmission following interspecies contact, extirpation probabilities for the seven bighorn sheep herds examined here ranged from 20% to 100%. The Disease Model allowed us to assess the probabilities that varied domestic sheep management scenarios would support persistent populations of free-ranging bighorn sheep.

Sexually antagonistic association between paternal phenotype and offspring viability reinforces total selection on a sexually selected trait.

The evolution of conspicuous sexually selected traits, such as horns or antlers, has fascinated biologists for more than a century. Elaborate traits can only evolve if they substantially increase reproduction, because they probably incur survival costs to the bearer. Total selection on these traits, however, includes sexual selection on sires and viability selection on offspring and can be influenced by changes in each of these components. Non-random associations between paternal phenotype and offspring viability may thus affect total selection on sexually selected traits. Long-term data on wild bighorn sheep (Ovis canadensis) provide the first evidence in nature that association between paternal phenotype and lamb viability strengthens total selection on horn size of adult rams, a sexually selected trait. The association of paternal horn length and offspring viability was sexually antagonistic: long-horned males sired sons with high viability but daughters of low viability. These results shed new light on the evolutionary dynamics of an iconic sexually selected trait and have important implications for sustainable wildlife management.