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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

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.

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.

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.