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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.

Migration is an effective behavioral strategy for prolonging access to seasonal resources and may be a resilient strategy for ungulates experiencing changing climatic conditions. In the Greater Yellowstone Ecosystem (GYE), elk are the primary ungulate, with approximately 20,000 individuals migrating to exploit seasonal gradients in forage while also avoiding energetically costly snow conditions. How climate-induced changes in plant phenology and snow accumulation are influencing elk migration timing is unknown. We present the most complete record of elk migration across the GYE, spanning 9 herds and 414 individuals from 2001 to 2017, to evaluate the drivers of migration timing and test for temporal shifts. The timing of elk departure from winter range involved a trade-off between current and anticipated forage conditions, while snow melt governed summer range arrival date. Timing of elk departure from summer range and arrival on winter range were both influenced by snow accumulation and exposure to hunting. At the GYE scale, spring and fall migration timing changed through time, most notably with winter range arrival dates becoming almost 50 days later since 2001. Predicted herd-level changes in migration timing largely agreed with observed GYE-wide changes-except for predicted winter range arrival dates which did not reflect the magnitude of change detected in the elk telemetry data. Snow melt, snow accumulation, and spring green-up dates all changed through time, with different herds experiencing different rates and directions of change. We conclude that elk migration is plastic, is a direct response to environmental cues, and that these environmental cues are not changing in a consistent manner across the GYE. The impacts of changing elk migration timing on predator-prey dynamics, carnivore-livestock conflict, disease ecology, and harvest management across the GYE are likely to be significant and complex.

1. While the tendency to return to previously visited locations - termed 'site fidelity' - is common in animals, the cause of this behaviour is not well understood. One hypothesis is that site fidelity is shaped by an animal's environment, such that animals living in landscapes with predictable resources have stronger site fidelity. Site fidelity may also be conditional on the success of animals' recent visits to that location, and it may become stronger with age as the animal accumulates experience in their landscape. Finally, differences between species, such as the way memory shapes site attractiveness, may interact with environmental drivers to modulate the strength of site fidelity.2. We compared inter-year site fidelity in 669 individuals across eight ungulate species fitted with GPS-collars and occupying a range of environmental conditions in North America and Africa. We used a distance-based index of site fidelity and tested hypothesized drivers of site fidelity using linear mixed effects models, while accounting for variation in annual range size.3. Mule deer Odocoileus hemionus and moose Alces alces exhibited relatively strong site fidelity, while wildebeest Connochaetes taurinus and barren-ground caribou Rangifer tarandus granti had relatively weak fidelity. Site fidelity was strongest in predictable landscapes where vegetative greening occurred at regular intervals over time (i.e. high temporal contingency). Species differed in their response to spatial heterogeneity in greenness (i.e. spatial constancy). Site fidelity varied seasonally in some species, but remained constant over time in others. Elk employed a 'win-stay, lose-switch' strategy, in which successful resource tracking in the springtime resulted in strong site fidelity the following spring. Site fidelity did not vary with age in any species tested.4. Our results provide support for the environmental hypothesis, particularly that regularity in vegetative phenology shapes the strength of site fidelity at the inter-annual scale. Large unexplained differences in site fidelity suggest that other factors, possibly species-specific differences in attraction to known sites, contribute to variation in the expression of this behaviour.5. Understanding drivers of variation in site fidelity across groups of organisms living in different environments provides important behavioural context for predicting how animals will respond to environmental change. published

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.

For ungulates and other long-lived species, life-history theory predicts that nutritional reserves are allocated to reproduction in a state-dependent manner because survival is highly conserved. Further, as per capita food abundance and nutritional reserves decline (i.e., density dependence intensifies), reproduction and recruitment become increasingly sensitive to weather. Thus, the degree to which weather influences vital rates should be associated with proximity to nutritional carrying capacity-a notion that we refer to as the Nutritional Buffer Hypothesis. We tested the Nutritional Buffer Hypothesis using six moose (Alces alces) populations that varied in calf recruitment (33-69 calves/100 cows). We predicted that populations with high calf recruitment were nutritionally buffered against the effects of unfavorable weather, and thus were below nutritional carrying capacity. We applied a suite of tools to quantify habitat and nutritional condition of each population and found that increased browse condition, forage quality, and body fat were associated with increased pregnancy and calf recruitment, thereby providing multiple lines of evidence that declines in calf recruitment were underpinned by resource limitation. From 2001 to 2015, recruitment was more sensitive to interannual variation in weather (e.g., winter severity, drought) and plant phenology (e.g., duration of spring) for populations with reduced browse condition, forage quality, and body fat, suggesting these populations lacked the nutritional reserves necessary to buffer demographic performance against the effects of unfavorable weather. Further, average within-population calf recruitment was determined by regional climatic variation, suggesting that the pattern of reduced recruitment near the southern range boundary of moose stems from an interaction between climate and resource limitation. When coupled with information on habitat, nutrition, weather, and climate, life-history theory provides a framework to estimate nutritional limitation, proximity to nutritional carrying capacity, and impacts of climate change for ungulates. Wyoming Game and Fish Department, Wyoming Wildlife Published version We thank Aimee Hurt, Ngaio Richards, Wicket, and Orbee from Working Dogs for Conservation for their assistance with locating moose feces, B. Davitt and the staff of the Washington State Wildlife Habitat Lab for estimating fecal N and NDF, J. Brown and the Center for Species Survival at the Smithsonian Conservation Biology Institute for quantifying fecal progestogen concentrations, J. Branen and the staff of BioTracking LLC for conducting the BioPryn Wild ELISA assays, the Matson Laboratory for analyzing tooth age, Jerod Merkle for assistance with quantifying plant phenology from NDVI data, and Melanie Murphy for providing laboratory space and guidance in fecal DNA analysis. This work was supported by grants from the Wyoming Game and Fish Department, Wyoming Wildlife Public domain – authored by a U.S. government employee

Ecological theory and empirical studies have demonstrated population-level demographic benefits resulting from a diversity of migratory behaviors with important implications for ecology, conservation, and evolution of migratory organisms. Nevertheless, evaluation of migratory portfolios (i.e., the variation in migratory behaviors across space and time among individuals within populations) has received relatively little attention in migratory ungulates, where research has focused largely on the dichotomous behaviors (e.g., resident and migrant) of partially migratory populations. Using GPS data from 361 female bighorn sheep (Ovis canadensis) across 17 (4 restored, 6 augmented, 7 native) populations in Montana and Wyoming, USA, we (1) characterized migratory portfolios based on behavioral and spatial migratory characteristics and (2) evaluated the relative influence of landscape attributes and management histories on migratory diversity. Native populations, which had been extant on the landscape for many generations, had more diverse migratory portfolios, higher behavioral switching rates, reduced seasonal range fidelity, and broad dispersion of individuals across summer and winter ranges. In contrast, restored populations with an abbreviated history on the landscape were largely non-migratory with a narrow portfolio of migratory behaviors, less behavioral switching, higher fidelity to seasonal ranges, and less dispersion on summer and winter ranges. Augmented populations were more variable and contained characteristics of both native and restored populations. Differences in migratory diversity among populations were associated with management histories (e.g., restored, augmented, or native). Landscape characteristics such as the duration and regularity of green-up, human landscape alterations, topography, and snow gradients were not strongly associated with migratory diversity. We suggest a two-pronged approach to restoring migratory portfolios in ungulates that first develops behavior-specific habitat models and then places individuals with known migratory behaviors into unoccupied areas in an effort to bolster migratory portfolios in restored populations, potentially with synergistic benefits associated with variation among individuals and resulting portfolio effects. Management efforts to restore diverse migratory portfolios may increase the abundance, resilience, and long-term viability of ungulate populations.

Landscape resistance is vital to connectivity modeling and frequently derived from resource selection functions (RSFs). RSFs estimate relative probability of use and tend to focus on understanding habitat preferences during slow, routine animal movements (e.g., foraging). Dispersal and migration, however, can produce rarer, faster movements, in which case models of movement speed rather than resource selection may be more realistic for identifying habitats that facilitate connectivity. To compare two connectivity modeling approaches applied to resistance estimated from models of movement rate and resource selection. Using movement data from migrating elk, we evaluated continuous time Markov chain (CTMC) and movement-based RSF models (i.e., step selection functions [SSFs]). We applied circuit theory and shortest random path (SRP) algorithms to CTMC, SSF and null (i.e., flat) resistance surfaces to predict corridors between elk seasonal ranges. We evaluated prediction accuracy by comparing model predictions to empirical elk movements. All connectivity models predicted elk movements well, but models applied to CTMC resistance were more accurate than models applied to SSF and null resistance. Circuit theory models were more accurate on average than SRP models. CTMC can be more realistic than SSFs for estimating resistance for fast movements, though SSFs may demonstrate some predictive ability when animals also move slowly through corridors (e.g., stopover use during migration). High null model accuracy suggests seasonal range data may also be critical for predicting direct migration routes. For animals that migrate or disperse across large landscapes, we recommend incorporating CTMC into the connectivity modeling toolkit.

Learning where and when Large ungulate migrations occur across continents and inspire curiosity about how these animals know when to leave and where to go. Jesmer et al. took advantage of regional extinctions and reintroductions of several North American ungulate species to determine the role of learning in migrations (see the Perspective by Festa-Bianchet). Reintroduced populations of bighorn sheep and moose did not migrate as historical herds had. However, after several decades, newly established herds were better able to track the emergence of vegetation in the environment and were increasingly migratory. Thus, newly introduced animals learned about their environment and shared the information through social exchange. Science , this issue p. 1023 ; see also p. 972

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.

Over the past two decades, populations of Shiras moose (Alces alces shirasi) in western Wyoming have declined. Recent work on the Jackson herd in northwest Wyoming suggests that the 1988 Yellowstone fires and regional drought contributed to a considerable decline in calf recruitment, which coincided with the recovery of grizzly bears (Ursus arctos horribilis) and gray wolves (Canis lupus occidentalis) to the Greater Yellowstone Ecosystem (GYE). Predation is also presumed to have contributed to declines in calf recruitment, but the relative influence of these predators has yet to be evaluated. We analyzed a time series of vital rates to identify the spatial extent and intensity at which predator density, habitat condition, and interacting abiotic factors limit population growth. We focused on neonate and winter calf survival because calves are the most vulnerable age class and because calf recruitment been shown to be one of the most important vital rates influencing population dynamics in large herbivores. Variables affecting calf survival were calculated at the seasonal home range scale from 100 individuals in the Jackson herd, and 90 individuals in an adjacent herd directly to the south, where moose are exposed to markedly lower predator density. We detected effects of grizzly bear density and habitat condition on neonate survival. Wolf density negatively influenced winter calf survival, but the effect was weak. Our spatial analysis of individual fitness in these two herds suggests that Shiras moose in the GYE have struggled to cope simultaneously with the effects of sub-optimal PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.2056v1 | CC-BY 4.0 Open Access | rec: 20 May 2016, publ: 20 May 2016 habitat conditions and predator density, highlighting the need to prioritize efforts to conserve Shiras moose as climate change continues to progress. PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.2056v1 | CC-BY 4.0 Open Access | rec: 20 May 2016, publ: 20 May 2016

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.

Many ungulate populations have lost access to their traditional migration routes and seasonal ranges, resulting in rapid and severe population declines. Some ungulate populations have been able to adapt to living year-round on one seasonal range and persist despite loss of migration. However, our understanding of how ungulates adapt their habitat selection and foraging strategies in order to compensate for migration loss is poor. This study investigates how a formerly migratory, now sedentary and isolated, bighorn sheep (Ovis canadensis) population persists year-round on high-elevation summer range in the Teton Range in northwest Wyoming. We captured and GPS-collared 20 bighorn ewes throughout the Teton Range in February 2008 and an additional 8 ewes in March 2009. In 2008, ninety percent of captured ewes were pregnant, and 100% were pregnant in 2009. During summers 2008 and 2009, we located and observed GPS-collared ewes, determined lamb survival, collected fecal samples for diet composition analysis, conducted vegetation surveys, and observed time-budgets. We found that 50% of lambs survived until at least mid-summer in 2008 and 60% survived in 2009. We observed differences in movement patterns between GPS-collared ewes during the summer seasons, ranging from 5 km to a maximum of 15 km. At this time, eight GPS-collared ewes have died (4 in avalanches, 1 from predation, and 3 unknown). This study is ongoing and will be completed in 2011. Results will directly contribute to management of this non-migratory and isolated bighorn sheep population, and will shed light on how a formerly migratory ungulate population has been able to persist on high-elevation range year-round

Many ungulate populations have lost access to their traditional migration routes and seasonal ranges, resulting in rapid and severe population declines. Some ungulate populations have been able to adapt to living year-round on one seasonal range and persist despite loss of migration. However, our understanding of how ungulates adapt their habitat selection and foraging strategies in order to compensate for migration loss is poor This study investigates how a formerly migratory, now sedentary and isolated bighorn sheep (Ovis canadensis) population persists year-round on high-elevation summer range in the Teton Range in northwest Wyoming. We captured and GPS-collared 20 bighorn ewes throughout the Teton Range in February 2008 and an additional 8 ewes in March 2009. In 2008, ninety percent of captured ewes were pregnant, and 100% were pregnant in 2009. During summer 2008, we located and observed GPS-collared ewes, determined lamb survival, collected fecal samples for diet composition analysis, and observed time-budgets. We found that 50% oflambs survived until at least mid­ summer. Diet composition analysis is pending and we are waiting to combine time-budget observations with additional data that will be collected during summers 2009 and 2010. We observed differences in movement patterns between GPS-collared ewes during summer 2008, ranging from 5 km to a maximum of 15 km. At this time, six GPS-collared ewes have died (four in avalanches, one from predation, and one unknown). This study is ongoing and will be completed in 2010. Results will directly contribute to management of this non-migratory and isolated bighorn sheep population, and will shed light on how a formerly migratory ungulate population has been able to persist on high­elevation range year-round.