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Abstract Many ungulates migrate between distinct summer and winter ranges, and identifying, mapping, and conserving these migration corridors have become a focus of local, regional, and global conservation efforts. Brownian bridge movement models (BBMMs) are commonly used to empirically identify these seasonal migration corridors; however, they require location data sampled at relatively frequent intervals to obtain a robust estimate of an animal's movement path. Fitting BBMMs to sparse location data violates the assumption of conditional random movement between successive locations, overestimating the area (and width) of a migration corridor when creating individual and population‐level occurrence distributions and precluding the use of low‐frequency, or sparse, data in mapping migration corridors. In an effort to expand the utility of BBMMs to include sparse GPS data, we propose an alternative approach to model migration corridors from sparse GPS data. We demonstrate this method using GPS data collected every 2 h from four mule deer (Odocoileus hemionus) and four elk (Cervus canadensis) herds within Wyoming and Idaho. First, we used BBMMs to estimate a baseline corridor for the 2‐h data. We then subsampled the 2‐h data to one location every 12 h (a proxy for sparse data) and fitted BBMMs to the 12‐h data using a fixed motion variance (FMV) value, instead of estimating the Brownian motion variance empirically. A range of FMV values was tested to identify the value that best approximated the baseline migration corridor. FMV values within a species‐specific range (mule deer: 400–1200 m2; elk: 600–1600 m2) successfully delineated migration corridors similar to the 2‐h baseline corridors; overall, lower values delineated narrower corridors and higher values delineated wider corridors. Optimal FMV values of 800 m2 (mule deer) and 1000 m2 (elk) decreased the inflation of the 12‐h corridors relative to the 2‐h corridors from traditional BBMMs. This FMV approach thus enables using sparse movement data to approximate realistic migration corridor dimensions, providing an important alternative when movement data are collected infrequently. This approach greatly expands the number of datasets that can be used for migration corridor mapping—a useful tool for management and conservation across the globe.

Abstract The spatial distribution of animals has consequences for nutrition, predator–prey dynamics, spread of diseases, and population dynamics in general. Animals must establish a home range to secure adequate resources to fuel their energy needs. Home ranges, therefore, are temporally and spatially dynamic, given the changing requirements of an animal and the availability of resources on the landscape. We used data from two populations of bighorn sheep with contrasting population dynamics following pneumonia epizootics and different habitat quality on their summer range to test the hypothesis that the distribution and size of home ranges are influenced by environmental conditions and reproductive status. We used a combination of data from 768 vegetation transects and remotely sensed metrics to index forage quality of consecutive biweekly home ranges for 27 bighorn sheep, June–August 2019–2021. There were population differences in home range dynamics that were consistent with resource limitations in the population declining in abundance. Animals in both populations increased the size of their home range through the summer in association with declining forage quality indexed by plant phenology. Furthermore, animals in the Whiskey Mountain population without offspring had home ranges more than twice the size of animals with offspring, whereas there were no differences in the home range size between animals with and without offspring in Jackson. We demonstrated that limitations young offspring impose on space use of a mother may have consequences for animals living where larger home ranges are needed to secure adequate resources—sheep on Whiskey Mountain had to travel 1000 m from escape terrain to access the same amount of biomass that the Jackson sheep could access directly adjacent to escape terrain. Forage quality and availability influence movement and space use. In the presence of disease, movement and space use may influence pathogen transmission and persistence. Thus, forage availability may play an indirect role in population dynamics in the presence of disease, which is another line of evidence for how environmental and nutritional conditions may influence population dynamics when coping with disease.

Abstract Observed links between parasites, such as ticks, and climate change have aroused concern for human health, wildlife population dynamics, and broader ecosystem effects. The one‐host life history of the winter tick (Dermacentor albipictus) links each annual cohort to environmental conditions during three specific time periods when they are predictably vulnerable: spring detachment from hosts, summer larval stage, and fall questing for hosts. We used mixed‐effects generalized linear models to investigate the drivers of tick loads carried by moose (Alces alces) relative to these time periods and across 750 moose, 10 years, and 16 study areas in the western United States. We tested for the effects of biotic factors (moose density, shared winter range, vegetation, migratory behavior) and weather conditions (temperature, snow, humidity) during each seasonal period when ticks are vulnerable and off‐host. We found that warm climatic regions, warm seasonal periods across multiple partitions of the annual tick life cycle, and warm years relative to long‐term averages each contributed to increased tick loads. We also found important effects of snow and other biotic factors such as host density and vegetation. Tick loads in the western United States were, on average, lower than those where tick‐related die‐offs in moose populations have occurred recently, but loads carried by some individuals may be sufficient to cause mortality. Lastly, we found interannual variation in tick loads to be most correlated with spring snowpack, suggesting this environmental component may have the highest potential to induce change in tick load dynamics in the immediate future of this region.

Abstract For various temperate ungulate species, recent research has highlighted the potential for spring vegetation phenology (“green‐up”) to influence individual condition, with purported benefits to population productivity. However, few studies have been able to measure the benefit on vital rates directly, and fewer still have investigated the comparative influence of other phenological periods on ungulate vital rates. In this study, we tracked phenological changes throughout the duration of the growing season and examined how their timing affected the probability of pregnancy in an ungulate population. We did this for elk (Cervus canadensis) across the Greater Yellowstone Ecosystem (GYE) by sampling 1106 adult females in winter at 25 sites over a 13‐year period and assessing sources of variation in pregnancy using a Bayesian hierarchical model. Pregnancy rates were generally high across the GYE (82.4%), and the primary influences on probability of pregnancy were the timing of vegetation senescence (“brown‐down”) in autumn and exposure to the reproductive disease brucellosis. Earlier forage brown‐down in fall negatively influenced the probability of pregnancy of elk aged 6–9 years by an estimated 17.2% within the range (ca. 32 days) of observed brown‐down end dates. While summer habitat quality has been inferred to influence elk pregnancy previously, our findings specify the key influence of foraging conditions later in the seasonal cycle, immediately before the breeding season. The reproductive disease brucellosis was also an important factor, reducing the probability of pregnancy by 12.4% in elk in the 6‐ to 9‐year age class. Because pregnancy was tested before most disease‐induced abortions occur, the apparent mechanism for this effect is a prolonged reduction in fertility beyond the period of initial exposure in which fetal mortality is typically expected. Our results prompt greater scrutiny of the combined effects of late‐season phenology and disease on reproductive rates and population productivity in temperate ungulates.

Abstract Food quality and availability, when combined with energetic demands in seasonal environments, shape resource acquisition and allocation by animals and hold consequences for life‐history strategies. In long‐lived species with extensive maternal care, regulation of somatic reserves of energy and protein can occur in a risk‐sensitive manner, wherein resources are preferentially allocated to support survival at the cost of investment in reproduction. We investigated how Rocky Mountain bighorn sheep (Ovis canadensis), an alpine mammal in a highly seasonal environment, allocates somatic reserves across seasons. In accordance with the hypothesis of risk‐sensitive resource allocation, we expected accretion and catabolism of somatic reserves to be regulated relative to preseason nutritional state, reproductive state, and variation among populations in accordance with local environmental conditions. To test that hypothesis, we monitored seasonal changes in percent ingesta‐free body fat (IFBFat) and ingesta‐free, fat‐free body mass (IFFFBMass) in three populations of bighorn sheep in northwest Wyoming between 2015 and 2019 through repeated captures of female sheep in December and March of each year in a longitudinal study design. Regulation of somatic reserves was risk‐sensitive and varied relative to the amount of somatic reserves an animal had at the beginning of the season. Regulation of fat reserves was sensitive to reproductive state and differed by population, particularly over the summer. In one population with low rates of recruitment of young, sheep that recruited offspring lost fat over the summer in contrast to the other two populations where sheep that recruited gained fat. And yet, all populations exhibited similar changes in fat catabolism and risk sensitivity over winter. The magnitude of body fat and mass change across seasons may be indicative of sufficiency of seasonal ranges to meet energetic demands of survival and reproduction. Risk‐sensitive allocation of resources was pervasive, suggesting nutritional underpinnings are foundational to behavior, vital rates, and, ultimately, population dynamics. For species living in alpine environments, risk‐sensitive resource allocation may be essential to balance investment in reproduction with ensuring survival.

Abstract Migration evolved as a behavior to enhance fitness through exploiting spatially and temporally variable resources and avoiding predation or other threats. Globally, landscape alterations have resulted in declines to migratory populations across taxa. Given the long time periods over which migrations evolved in native systems, it is unlikely that restored populations embody the same migratory complexity that existed before population reductions or regional extirpation. We used GPS location data collected from 209 female bighorn sheep (Ovis canadensis) to characterize population and individual migration patterns along elevation and geographic continuums for 18 populations of bighorn sheep with different management histories (i.e., restored, augmented, and native) across the western United States. Individuals with resident behaviors were present in all management histories. Elevational migrations were the most common population‐level migratory behavior. There were notable differences in the degree of individual variation within a population across the three management histories. Relative to native populations, restored and augmented populations had less variation among individuals with respect to elevation and geographic migration distances. Differences in migratory behavior were most pronounced for geographic distances, where the majority of native populations had a range of variation that was 2–4 times greater than restored or augmented populations. Synthesis and applications. Migrations within native populations include a variety of patterns that translocation efforts have not been able to fully recreate within restored and augmented populations. Theoretical and empirical research has highlighted the benefits of migratory diversity in promoting resilience and population stability. Limited migratory diversity may serve as an additional factor limiting demographic performance and range expansion. We suggest preserving native systems with intact migratory portfolios and a more nuanced approach to restoration and augmentation in which source populations are identified based on a suite of criteria that includes matching migratory patterns of source populations with local landscape attributes.

Abstract In the Rocky Mountains, bighorn sheep restoration has been only marginally effective; this iconic wilderness species currently exists at a fraction of their historic abundance and often in fragmented and small populations. To inform bighorn sheep conservation and restoration efforts, it is critical to understand sources of variation in key vital rates. Our objectives were to characterize the spatiotemporal variations and factors affecting survival and pregnancy rates of bighorn sheep (Ovis canadensis) using data from 19 bighorn sheep populations in Montana and Wyoming that occupied diverse landscapes ranging from the Northern Great Plains to the Rocky Mountains. We used a hierarchical modeling approach to estimate survival and pregnancy rates of adult females and identify the important intrinsic and environmental factors affecting these vital rates. Survival of prime‐aged animals was relatively high and stable, and pregnancy rates for prime‐aged animals showed more overall variation in response to intrinsic and extrinsic factors. Summer growing season, as indexed by integrated NDVI, positively influenced the probability of pregnancy and winter survival. This highlights the important relationship between summer growing season conditions and bighorn sheep physiological status. An index of mountain lion population abundance was related weakly to winter survival of bighorn sheep, with mountain lion abundance on winter ranges negatively affecting winter survival. Our results regarding the distribution of the estimated probabilities of pregnancy and survival, and the identification of factors associated with regional variability in these vital rates provide a foundation for understanding the dynamics of bighorn sheep populations in the Rocky Mountains. The importance of summer growing season conditions suggests management efforts should focus on maintaining and improving nutritional resources on bighorn sheep summer ranges, in efforts to enhance the condition of animals entering the breeding season and nutrient‐limited winter season. Although we document nontrivial changes in both pregnancy and winter survival rates associated with environmental variation, our results broadly support the dominant paradigm of ungulate demography insofar as survival rates of adult females were relatively higher and had less variation than pregnancy rates.

ABSTRACT Population growth and range expansion of nonnative species can potentially disrupt ecosystem function or add conservation value to an area, and evaluation of possible impacts can be a challenge for managers. Nonnative populations of mountain goats (Oreamnos americanus) are present in the Greater Yellowstone Area (GYA) in the U.S. states of Idaho, Montana, and Wyoming because of historical introduction events, but their population trend and range have not been assessed across the area. We used 6,701 location records from 1947 to 2015 to map mountain goat distribution and evaluate, in a descriptive manner, range overlap with native bighorn sheep (Ovis canadensis). We analyzed 136 survey counts using the Exponential Growth State–Space model to estimate population trends and abundance. Mountain goats dispersed 50–85 km from introduction sites to occupy all mountain ranges in the northern GYA and 30–40 km to occupy new areas in the southern GYA. Mountain goat numbers increased in nearly all count units, with the strongest growth rates estimated in areas more recently colonized. Using moderate detection probability (0.70), we estimated approximately 2,355 mountain goats in the GYA. Although not tested in our analysis, the gradual range expansion and population growth rates were consistent with density‐dependent processes observed in other introduced large herbivores and demonstrate that mountain goats can successfully disperse over unsuitable locales to colonize new areas. Therefore, we expect mountain goat populations will continue to expand into unoccupied mountain ranges that contain significant numbers of bighorn sheep unless specific management actions are implemented to address their population growth. © 2016 The Wildlife Society.

Abstract The persistence of many migratory ungulate populations worldwide is threatened due to anthropogenic impacts to seasonal ranges and migration routes. While many studies have linked migratory ungulate declines to migration disruption or loss, very few have explored the underlying factors that determine whether a population perishes or persists. In some cases, populations undergo severe declines and extirpation after migration loss; however, others appear able to persist as residents. We predict that to persist, populations must replace the traditional benefits of migration by altering the foraging strategies they employ as residents within one seasonal range. We propose the alternative foraging strategies (AFS) hypothesis as a framework for identifying various behavioral strategies that populations may use to cope with migration loss. We tested the hypothesis using the formerly migratory Teton bighorn sheep population in northwest Wyoming, which ceased migrating over 60 yr ago, but has persisted as a resident population. We used global positioning system data to evaluate winter and summer habitat selection and seasonal elevational movements for 28 adult female bighorn sheep (Ovis canadensis) from 2008 to 2010. Resource selection functions revealed that bighorn sheep employ winter foraging strategies to survive as residents by seeking out rugged, high‐elevation, windswept ridgelines. Seasonal movement analyses indicated that bighorn sheep undergo a newly documented “abbreviated migration” strategy that is closely synchronized with vegetation green‐up patterns within their one range. Bighorn sheep descend 500 m in elevation and travel up to 10 km in spring, gaining access to newly emergent forage approximately 30 d before it appears on their high‐elevation winter and summer ranges. Our findings indicate that the Teton bighorn sheep population has persisted due to its habitat selection, AFS, and unique movement patterns, which allow migration loss to be mediated to some extent. The identification of AFS and the habitats that support them can help reveal the underlying benefits of migration and conserve populations in the face of future migration loss.

Respiratory disease caused by Mycoplasma ovipneumoniae and Pasteurellaceae poses a formidable challenge for bighorn sheep (Ovis canadensis) conservation. All-age epizootics can cause 10-90% mortality and are typically followed by multiple years of enzootic disease in lambs that hinders post-epizootic recovery of populations. The relative frequencies at which these epizootics are caused by the introduction of novel pathogens or expression of historic pathogens that have become resident in the populations is unknown. Our primary objectives were to determine how commonly the pathogens associated with respiratory disease are hosted by bighorn sheep populations and assess demographic characteristics of populations with respect to the presence of different pathogens. We sampled 22 bighorn sheep populations across Montana and Wyoming, USA for Mycoplasma ovipneumoniae and Pasteurellaceae and used data from management agencies to characterize the disease history and demographics of these populations. We tested for associations between lamb:ewe ratios and the presence of different respiratory pathogen species. All study populations hosted Pasteurellaceae and 17 (77%) hosted Mycoplasma ovipneumoniae. Average lamb:ewe ratios for individual populations where both Mycoplasma ovipneumoniae and Pasteurellaceae were detected ranged from 0.14 to 0.40. However, average lamb:ewe ratios were higher in populations where Mycoplasma ovipneumoniae was not detected (0.37, 95% CI: 0.27-0.51) than in populations where it was detected (0.25, 95% CI: 0.21-0.30). These findings suggest that respiratory pathogens are commonly hosted by bighorn sheep populations and often reduce recruitment rates; however ecological factors may interact with the pathogens to determine population-level effects. Elucidation of such factors could provide insights for management approaches that alleviate the effects of respiratory pathogens in bighorn sheep. Nevertheless, minimizing the introduction of novel pathogens from domestic sheep and goats remains imperative to bighorn sheep conservation.