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The ability to freely move across the landscape to track the emergence of nutritious spring green-up (termed 'green-wave surfing') is key to the foraging strategy of migratory ungulates. Across the vast landscapes traversed by many migratory herds, habitats are being altered by development with unknown consequences for surfing. Using a unique long-term tracking dataset, we found that when energy development occurs within mule deer (Odocoileus hemionus) migration corridors, migrating animals become decoupled from the green wave. During the early phases of a coalbed natural gas development, deer synchronized their movements with peak green-up. But faced with increasing disturbance as development expanded, deer altered their movements by holding up at the edge of the gas field and letting the green wave pass them by. Development often modified only a small portion of the migration corridor but had far-reaching effects on behaviour before and after migrating deer encountered it, thus reducing surfing along the entire route by 38.65% over the 14-year study period. Our study suggests that industrial development within migratory corridors can change the behaviour of migrating ungulates and diminish the benefits of migration. Such disruptions to migratory behaviour present a common mechanism whereby corridors become unprofitable and could ultimately be lost on highly developed landscapes.

Aim: Macroecological studies that require habitat suitability data for many species often derive this information from expert opinion. However, expert- based informa-tion is inherently subjective and thus prone to errors. The increasing availability of GPS tracking data offers opportunities to evaluate and supplement expert- based in-formation with detailed empirical evidence. Here, we compared expert- based habitat suitability information from the International Union for Conservation of Nature (IUCN) with habitat suitability information derived from GPS-tracking data of 1,498 individu-als from 49 mammal species.Location: Worldwide.Time period: 1998–2021.Major taxa studied: Forty- nine terrestrial mammal species.Methods: Using GPS data, we estimated two measures of habitat suitability for each individual animal: proportional habitat use (proportion of GPS locations within a habi-tat type), and selection ratio (habitat use relative to its availability). For each individual we then evaluated whether the GPS- based habitat suitability measures were in agree-ment with the IUCN data. To that end, we calculated the probability that the ranking of empirical habitat suitability measures was in agreement with IUCN's classification into suitable, marginal and unsuitable habitat types.Results: IUCN habitat suitability data were in accordance with the GPS data (> 95% probability of agreement) for 33 out of 49 species based on proportional habitat use estimates and for 25 out of 49 species based on selection ratios. In addition, 37 and 34 species had a > 50% probability of agreement based on proportional habitat use and selection ratios, respectively.Main conclusions: We show how GPS-tracking data can be used to evaluate IUCN habitat suitability data. Our findings indicate that for the majority of species in-cluded in this study, it is appropriate to use IUCN habitat suitability data in macro-ecological studies. Furthermore, we show that GPS-tracking data can be used to identify and prioritize species and habitat types for re-evaluation of IUCN habitat suitability data. expert opinion, GPS, habitat suitability, habitat type, habitat use, IUCN, mammals, movement, selection ratio, telemetry

Our understanding of ungulate migration is advancing rapidly due to innovations in modern animal tracking. Herein, we review and synthesize nearly seven decades of work on migration and other long-distance movements of wild ungulates. Although it has long been appreciated that ungulates migrate to enhance access to forage, recent contributions demonstrate that their movements are fine tuned to dynamic landscapes where forage, snow, and drought change seasonally. Researchers are beginning to understand how ungulates navigate migrations, with the emerging view that animals blend gradient tracking with spatial memory, some of which is socially learned. Although migration often promotes abundant populations—with broad effects on ecosystems—many migrations around the world have been lost or are currently threatened by habitat fragmentation, climate change, and barriers to movement. Fortunately, new efforts that use empirical tracking data to map migrations in detail are facilitating effective conservation measures to maintain ungulate migration.

Fences have recently been recognized as one of the most prominent linear infrastructures on earth. As animals traverse fenced landscapes, they adjust movement behaviors to optimize resource access while minimizing energetic costs of coping with fences. Examining individual responses is key for connecting localized fence effects with population dynamics. We investigated the multi-scale effects of fencing on animal movements, space use, and survival of 61 pronghorn and 96 mule deer on a gradient of fence density in Wyoming, USA. Taking advantage of the recently developed Barrier Behavior Analysis, we classified individual movement responses upon encountering fences (i.e. barrier behaviors). We adopted the reaction norm framework to jointly quantify individual plasticity and behavioral types of barrier behaviors, as well as behavior syndromes between barrier behaviors and animal space use. We also assessed whether barrier behaviors affect individual survival. Our results highlighted a high level individual plasticity encompassing differences in the degree and direction of barrier behaviors for both pronghorn and mule deer. Additionally, these individual differences were greater at higher fence densities. For mule deer, fence density determined the correlation between barrier behaviors and space use, and was negatively associated with individual survival. Yet, these relationships were not statistically significant for pronghorn. By integrating approaches from movement ecology and behavioral ecology with the emerging field of fence ecology, this study provides new evidence that an extraordinarily widespread linear infrastructure uniquely impacts animals at the individual level. Managing landscape for lower fence densities may help prevent irreversible behavioral shifts for wide-ranging animals in fenced landscapes.

While migrating, animals make directionally persistent movements and may only respond to human-induced rapid environmental change (HIREC), such as climate and land-use change, once a threshold of HIREC is surpassed. In contrast, animals on other seasonal ranges (e.g., winter range) make more localized and tortuous movements while foraging and may have the flexibility to adjust the location of their range and the intensity of use within it to minimize interactions with HIREC. Because of these seasonal differences in movement, animals on seasonal ranges should avoid areas that contain any level of HIREC, however, during migration, animals should use areas that contain low levels of HIREC, avoiding it only once a threshold of HIREC has been surpassed. We tested this hypothesis using a decade of GPS collar data collected from migratory mule deer (Odocoileus hemionus; n = 56 migration, 143 winter) and pronghorn (Antilocapra americana; n = 70 migration, 89 winter) that winter on and migrate through a natural gas field in western Wyoming. Using surface disturbance caused by well pads and roads as an index of HIREC, we evaluated behavioral responses across three spatial scales during winter and migration seasons. During migration, both species tolerated low levels of disturbance. Once a disturbance threshold was surpassed, however, they avoided HIREC. For mule deer, thresholds were consistently ~3%, whereas thresholds for pronghorn ranged from 1% to 9.25% surface disturbance. In contrast to migration, both species generally avoided all levels of HIREC while on winter range. Our study suggests that animal responses to HIREC are mediated by season-specific movement patterns. Our results provide further evidence of ungulates avoiding human disturbance on winter range and reveal disturbance thresholds that trigger mule deer and pronghorn responses during migration: information that managers can use to maintain the ecological function of migration routes and winter ranges.

Maintaining functional connectivity is critical for the long‐term conservation of wildlife populations. Landscape genomics provides an opportunity to assess long‐term functional connectivity by relating environmental variables to spatial patterns of genomic variation resulting from generations of movement, dispersal and mating behaviors. Identifying landscape features associated with gene flow at large geographic scales for highly mobile species is becoming increasingly possible due to more accessible genomic approaches, improved analytical methods and enhanced computational power. We characterized the genetic structure and diversity of migratory mule deer Odocoileus hemionus using 4051 single nucleotide polymorphisms in 406 individuals sampled across multiple habitats throughout Wyoming, USA. We then identified environmental variables associated with genomic variation within genetic groups and statewide using a stepwise approach to first evaluate nonlinear relationships of landscape resistance with genetic distances and then use mixed‐effects modeling to choose top landscape genomic models. We identified three admixed genetic groups of mule deer and found that environmental variables associated with gene flow varied among genetic groups, revealing scale‐dependent and regional variation in functional connectivity. At the statewide scale, more gene flow occurred in areas with low elevation and mixed habitat. In the southern genetic group, more gene flow occurred in areas with low elevation. In the northern genetic group, more gene flow occurred in grassland and forest habitats, while highways and energy infrastructure reduced gene flow. In the western genetic group, the null model of isolation by distance best represented genetic patterns. Overall, our findings highlight the role of different seasonal ranges on mule deer genetic connectivity, and show that anthropogenic features hinder connectivity. This study demonstrates the value of combining a large, genome‐wide marker set with recent advances in landscape genomics to evaluate functional connectivity in a wide‐ranging migratory species.

Migration of ungulates (hooved mammals) is a fundamental ecological process that promotes abundant herds, whose effects cascade up and down terrestrial food webs. Migratory ungulates provide the prey base that maintains large carnivore and scavenger populations and underpins terrestrial biodiversity (fig. S1). When ungulates move in large aggregations, their hooves, feces, and urine create conditions that facilitate distinct biotic communities. The migrations of ungulates have sustained humans for thousands of years, forming tight cultural links among Indigenous people and local communities. Yet ungulate migrations are disappearing at an alarming rate (1). Efforts by wildlife managers and conservationists are thwarted by a singular challenge: Most ungulate migrations have never been mapped in sufficient detail to guide effective conservation. Without a strategic and collaborative effort, many of the world's great migrations will continue to be truncated, severed, or lost in the coming decades. Fortunately, a combination of animal tracking datasets, historical records, and local and Indigenous knowledge can form the basis for a global atlas of migrations, designed to support conservation action and policy at local, national, and international levels.

The forage maturation hypothesis (FMH) states that energy intake for ungulates is maximised when forage biomass is at intermediate levels. Nevertheless, metabolic allometry and different digestive systems suggest that resource selection should vary across ungulate species. By combining GPS relocations with remotely sensed data on forage characteristics and surface water, we quantified the effect of body size and digestive system in determining movements of 30 populations of hindgut fermenters (equids) and ruminants across biomes. Selection for intermediate forage biomass was negatively related to body size, regardless of digestive system. Selection for proximity to surface water was stronger for equids relative to ruminants, regardless of body size. To be more generalisable, we suggest that the FMH explicitly incorporate contingencies in body size and digestive system, with small-bodied ruminants selecting more strongly for potential energy intake, and hindgut fermenters selecting more strongly for surface water.

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

Long-distance migration by terrestrial mammals is a phenomenon critical to the persistence of populations, but such migrations are declining globally because of over-harvest, habitat loss, and movement barriers. Increasingly, there is a need to improve existing routes, mitigate route segments affected by anthropogenic disturbance, and in some instances, determine whether alternative routes are available. Using a hypothesis-driven approach, we identified landscape features associated with the primary functional attributes, stopovers and movement corridors, of spring migratory routes for mule deer in two study areas using resource selection functions. Patterns of selection for landscape attributes of movement corridors and stopovers mostly were similar; however, landscape features associated with movement corridors aligned better with areas that facilitated movement, whereas selection of stopovers was consistent with sites offering early access to spring forage. For movement corridors, deer selected for dry sites, low elevation, and low anthropogenic disturbance. For stopovers, deer selected for dry sites, with consistently early green-up across years, south-southwesterly aspects, low elevation, and low anthropogenic disturbance. Stopovers and movement corridors of a migratory route presumably promote different functions, but for a terrestrial migrant, patterns of habitat selection indicate that the same general habitat attributes may facilitate both movement and foraging in spring. Our findings emphasize the roles of topographical wetness, vegetation phenology, and anthropogenic disturbance in shaping use of the landscape during migration for this large herbivore. Avoiding human disturbance and tracking ephemeral forage resources appear to be a consistent pattern during migration, which reinforces the notion that movement during migration has a nutritional underpinning and disturbance potentially alters the net benefits of migration.

Summary Animals exhibit a diversity of movement tactics [ 1 ]. Tracking resources that change across space and time is predicted to be a fundamental driver of animal movement [ 2 ]. For example, some migratory ungulates (i.e., hooved mammals) closely track the progression of highly nutritious plant green-up, a phenomenon called “green-wave surfing” [ 3 , 4 , 5 ]. Yet general principles describing how the dynamic nature of resources determine movement tactics are lacking [ 6 ]. We tested an emerging theory that predicts surfing and the existence of migratory behavior will be favored in environments where green-up is fleeting and moves sequentially across large landscapes (i.e., wave-like green-up) [ 7 ]. Landscapes exhibiting wave-like patterns of green-up facilitated surfing and explained the existence of migratory behavior across 61 populations of four ungulate species on two continents (n = 1,696 individuals). At the species level, foraging benefits were equivalent between tactics, suggesting that each movement tactic is fine-tuned to local patterns of plant phenology. For decades, ecologists have sought to understand how animals move to select habitat, commonly defining habitat as a set of static patches [ 8 , 9 ]. Our findings indicate that animal movement tactics emerge as a function of the flux of resources across space and time, underscoring the need to redefine habitat to include its dynamic attributes. As global habitats continue to be modified by anthropogenic disturbance and climate change [ 10 ], our synthesis provides a generalizable framework to understand how animal movement will be influenced by altered patterns of resource phenology.

An increasing global demand for energy assures continued disturbance to previously undeveloped landscapes, but understanding broader impacts to wildlife remains elusive. Among groups of species most vulnerable to habitat disruption are those requiring large tracts of land. Pronghorn (Antilocapra americana) are an obligate to the open plains and basins that, similar to other transcontinental large herbivores, rely primarily on habitats where development of energy resources such as oil, natural gas, coal, wind, and solar are intensifying. To understand behavioral response to a burgeoning energy development project, we evaluated avoidance, displacement, and winter residency patterns of pronghorn in the southern Greater Yellowstone Ecosystem across a 15‐year period using 171 collared individuals. Distance from natural gas well pads increased through time and was concurrent with declines in winter residency. Between 2005 and 2017, we found that (a) pronghorn avoidance of well pads likely increased by 408 m, (b) the overall displacement of pronghorn relative to well pads in the final year of study increased by 800 m, (c) the time pronghorn spent in the study area was reduced by 22% (nearly 1 month), and (d) the percentage of pronghorn leaving the study area increased by 57%. Such directional changes signal a strong behavioral response of an open‐plain obligate to energy infrastructure, and together, these metrics indicate that pronghorn response to energy development involves both avoidance of infrastructure and partial abandonment of their traditional winter ranges. While comparable long‐term data sets are generally unavailable for other functionally equivalent ungulate groups in similar ecological topographies of Asia, Africa, and South America, our study may serve as a reasonable surrogate and highlights that behavioral changes elicited from energy development which at first appear subtle can proliferate and may portend demographic consequences.

Our knowledge of migration ecology has progressed quickly in concert with technological advances that collect fine‐scale movement data through time. We now know that migration plays a critical role in the annual nutritional cycle of large herbivores and that sustaining functional migratory routes is key to long‐term conservation. Yet, we lack basic information on whether one migratory route may function better than another, or more specifically, if choosing one route over another has fitness consequences – knowledge that could help inform conservation and restoration efforts. Here, we examined how a suite of migratory parameters influenced the survival of mule deer Odocoileus hemionus that shared a common winter range in New Mexico, USA, but migrated to various summer ranges in Colorado, USA. We used a Cox proportional hazard model and longitudinal global positioning system data collected over a 7‐year period to investigate whether the mortality risk of 66 deer was affected by choice of migratory route, summer range, migration distance, speed or the number of administrative boundaries each route crossed. We found mule deer survival was not influenced by migratory distance, speed or number of administrative boundaries, but was strongly affected by the choice of migratory route and summer range. The magnitude of these effects was surprisingly large, doubling or tripling mortality risk. Cumulative survival rates showed that regardless of summer range, individuals migrating along high‐use exterior routes had cumulative survival rates approximately 30% lower than individuals migrating along high‐use interior routes. To our knowledge, this is the first direct evidence that a mammal's choice of migration route can influence its probability of survival. Synthesis and applications. Our finding that large herbivores may experience up to three times higher mortality risk by using a different migratory route reveals a novel link between migration and demography. Importantly, our results also suggest that spatially explicit model parameters and predictions could help in the conservation and restoration of migratory populations by identifying specific migratory routes or seasonal ranges that reduce survival.

From fine-scale foraging to broad-scale migration, animal movement is shaped by the distribution of resources. There is mounting evidence, however, that learning and memory also guide movement. Although migratory mammals commonly track resource waves, how resource tracking and memory guide long-distance migration has not been reconciled. We examined these hypotheses using movement data from four populations of migratory mule deer (n = 91). Spatial memory had an extraordinary influence on migration, affecting movement 2-28 times more strongly than tracking spring green-up or autumn snow depth. Importantly, with only an ability to track resources, simulated deer were unable to recreate empirical migratory routes. In contrast, simulated deer with memory of empirical routes used those routes and obtained higher foraging benefits. For migratory terrestrial mammals, spatial memory provides knowledge of where seasonal ranges and migratory routes exist, whereas resource tracking determines when to beneficially move within those areas.

The migratory movements of wild animals can promote abundance and support ecosystem functioning. For large herbivores, mounting evidence suggests that migratory behaviour is an individually variable trait, where individuals can easily switch between migrant and resident tactics. The degree of migratory plasticity, including whether and where to migrate, has important implications for the ecology and conservation of large herbivores in a changing world. Mule deer (Odocoileus hemionus) are an iconic species of western North America, but are notably absent from the body of literature that suggests large herbivore migrations are highly plastic. We evaluated plasticity of migration in female mule deer using longitudinal GPS data collected from 312 individuals across nine populations in the western United States, including 882 animal-years (801 migrants and 81 residents). We followed both resident and migratory mule deer through time to determine whether individual animals switched migratory behaviours (i.e., whether to migrate) from migratory to residency or vice versa. Additionally, we examined the fidelity of individuals to their migration routes (i.e., where to migrate) to determine whether they used the same routes year after year. We also evaluated whether age and reproductive status affected propensity to migrate or fidelity to migratory routes. Our results indicate that mule deer, unlike other large herbivores, have little or no plasticity in terms of whether or where they migrate. Resident deer remained residents, and migrant deer remained migrants, regardless of age, reproductive status or number of years monitored. Further, migratory individuals showed strong fidelity (>80%) to their migration routes year after year. Our study clearly shows that migration plasticity is not ubiquitous among large herbivores. Because of their rigid migratory behaviour, mule deer may not adapt to changing environmental conditions as readily as large herbivores with more plastic migratory behaviour (e.g., elk). The fixed migratory behaviours of mule deer make clear that conservation efforts aimed at traditional seasonal ranges and migration routes are warranted for sustaining this iconic species that continues to decline across its range.

The availability and quality of forage on the landscape constitute the foodscape within which animals make behavioral decisions to acquire food. Novel changes to the foodscape, such as human disturbance, can alter behavioral decisions that favor avoidance of perceived risk over food acquisition. Although behavioral changes and population declines often coincide with the introduction of human disturbance, the link(s) between behavior and population trajectory are difficult to elucidate. To identify a pathway by which human disturbance may affect ungulate populations, we tested the Behaviorally Mediated Forage‐Loss Hypothesis, wherein behavioral avoidance is predicted to reduce use of available forage adjacent to disturbance. We used GPS collar data collected from migratory mule deer (Odocoileus hemionus) to evaluate habitat selection, movement patterns, and time‐budgeting behavior in response to varying levels of forage availability and human disturbance in three different populations exposed to a gradient of energy development. Subsequently, we linked animal behavior with measured use of forage relative to human disturbance, forage availability, and quality. Mule deer avoided human disturbance at both home range and winter range scales, but showed negligible differences in vigilance rates at the site level. Use of the primary winter forage, sagebrush (Artemisia tridentata), increased as production of new annual growth increased but use decreased with proximity to disturbance. Consequently, avoidance of human disturbance prompted loss of otherwise available forage, resulting in indirect habitat loss that was 4.6‐times greater than direct habitat loss from roads, well pads, and other infrastructure. The multiplicative effects of indirect habitat loss, as mediated by behavior, impaired use of the foodscape by reducing the amount of available forage for mule deer, a consequence of which may be winter ranges that support fewer animals than they did before development.

Rapid climate and human land-use change may limit the ability of long-distance migratory herbivores to optimally track or ‘surf’ high-quality forage during spring green-up. Understanding how anthropogenic and environmental stressors influence migratory movements is of critical importance because of their potential to cause a mismatch between the timing of animal movements and the emergence of high-quality forage. We measured stress hormones (fecal glucocorticoid metabolites; FGMs) to test hypotheses about the effects of high-quality forage tracking, human land-use and use of stopover sites on the physiological state of individuals along a migratory route. We collected and analysed FGM concentrations from 399 mule deer (Odocoileus hemionus) samples obtained along a 241-km migratory route in western Wyoming, USA, during spring 2015 and 2016. In support of a fitness benefit hypothesis, individuals occupying areas closer to peak forage quality had decreased FGM levels. Specifically, for every 10-day interval closer to peak forage quality, we observed a 7% decrease in FGMs. Additionally, we observed support for both an additive anthropogenic stress hypothesis and a hypothesis that stopovers act as physiological refugia, wherein individuals sampled far from stopover sites exhibited 341% higher FGM levels if in areas of low landscape integrity compared to areas of high landscape integrity. Overall, our findings indicate that the physiological state of mule deer during migration is influenced by both anthropogenic disturbances and their ability to track high-quality forage. The availability of stopovers, however, modulates physiological responses to those stressors. Thus, our results support a recent call for the prioritization of stopover locations and connectivity between those locations in conservation planning for migratory large herbivores.

The seasonal migrations of ungulates are increasingly threatened by various forms of anthropogenic disturbance, including roads, fences, and other infrastructure. Although roadway impacts (e.g., wildlife–vehicle collisions and landscape permeability) to species such as mule deer (Odocoileus hemionus) can largely be mitigated with underpasses and continuous fencing, similar mitigation may not be effective for pronghorn (Antilocapra americana) or other ungulate species that are reluctant to move through confined areas. The Wyoming Department of Transportation recently installed 6 underpasses and 2 overpasses along 20 km of U.S. Highway 191 in western Wyoming, USA, where we evaluated species-specific preferences by documenting the number of migratory mule deer and pronghorn that used adjacent overpasses and underpasses for 3 years (autumn 2012 to spring 2015) following construction. We also measured the amount of back-and-forth movement across the highway for each species through time. We documented 40,251 crossings of the highway by mule deer and 19,290 crossings by pronghorn. Of those highway crossings, 79% of mule deer moved under, whereas 93% of pronghorn moved over the highway. These strong species-specific differences were evident at both sites and support the notion that overpasses are more amenable to pronghorn than underpasses. Concurrently, we documented a 64–306% increase in the amount of back-and-forth movement of mule deer and pronghorn across the highway during migration periods. Such movement flexibility is presumed to improve their ability to respond to changing environmental conditions by easily accessing habitats on either side of the highway. Our results highlight that species-specific preferences are an important consideration when mitigating roadway impacts with wildlife crossing structures. © 2016 The Wildlife Society.

Made available in DSpace on 2018-11-26T17:44:52Z (GMT). No. of bitstreams: 0 Previous issue date: 2018-01-26 Robert Bosch Foundation Animal movement is fundamental for ecosystem functioning and species survival, yet the effects of the anthropogenic footprint on animal movements have not been estimated across species. Using a unique GPS-tracking database of 803 individuals across 57 species, we found that movements of mammals in areas with a comparatively high human footprint were on average one-half to one-third the extent of their movements in areas with a low human footprint. We attribute this reduction to behavioral changes of individual animals and to the exclusion of species with long-range movements from areas with higher human impact. Global loss of vagility alters a key ecological trait of animals that affects not only population persistence but also ecosystem processes such as predator-prey interactions, nutrient cycling, and disease transmission. 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WY USA North Carolina Museum Nat Sci, Raleigh, NC 27601 USA North Carolina State Univ, Dept Forestry & Environm Resources, Raleigh, NC 27695 USA Karatina Univ, Dept Nat Resource Management, POB 195710101, Karatina, Kenya Univ Lethbridge, Dept Psychol, Lethbridge, AB T1K 3M4, Canada Columbia Univ, Lamont Doherty Earth Observ, Palisades, NY 10964 USA Univ Valencia, Terr Vertebrates Grp, Cavanilles Inst Biodivers & Evolutionary Biol, E-46980 Valencia, Spain SUNY Stony Brook, Dept Anthropol, Stony Brook, NY 11794 USA TSG, SSC, IUCN, Rua Licuala 622,Damha 1,Campo Grande, BR-79046150 Mato Grosso Sul, Brazil IPE Inst Pesquisas Ecol Inst Ecol Res, Caixa Postal 47, BR-12960000 Sao Paulo, Brazil Univ Alicante, Dept Ciencias Ambient & Recursos Natur, Vertebrates Zool Res Grp, Alicante, Spain Embrapa Pantanal, BR-79320900 Corumba, MS, Brazil Chico Mendes Inst Conservat Biodivers, Natl Res Ctr Carnivores Conservat, BR-12952011 Atiba, SP, Brazil Univ Glasgow, Inst Biodivers Anim Hlth & Comparat Med, Glasgow, Lanark, Scotland NYU, Dept Biol, Ctr Genom & Syst Biol, New York, NY 10003 USA Univ Calif Davis, Dept Microbiol & Mol Genet, Davis, CA 95616 USA Univ Oslo, Dept Biosci, Ctr Ecol & Evolutionary Synth, NO-0316 Oslo, Norway Hebrew Univ Jerusalem, Alexander Silberman Inst Life Sci, Dept Ecol Evolut & Behav, Movement Ecol Lab, IL-91904 Jerusalem, Israel Norwegian Inst Nat Res, NO-0349 Oslo, Norway Norwegian Univ Sci & Technol NTNU, Dept Math Sci & Ctr Biodivers Dynam, N-7491 Trondheim, Norway Univ Fed Mato Grosso do Sul, Dept Ecol, BR-79070900 Campo Grande, MS, Brazil Field Museum Nat Hist, Integrat Res Ctr, Chicago, IL 60605 USA Consorzio Parco Nazl Stelvio, Bormio, Sondrio, Italy Univ Grenoble Alpes, Irstea, UR LESSEM, BP 76, F-38402 St Martin Dheres, France Univ Bayreuth, BayCEER, D-95447 Bayreuth, Germany Natl Park Schwarzwald, D-77889 Seebach, Germany Univ New South Wales, Evolut & Ecol Res Ctr, Sydney, NSW 2052, Australia Univ New South Wales, Sch Biol Earth & Environm Sci, Sydney, NSW 2052, Australia Penn Game Commiss, Harrisburg, PA 17110 USA Princeton Univ, Dept Ecol & Evolutionary Biol, Princeton, NJ 08544 USA Univ Konstanz, Dept Biol, D-78467 Constance, Germany Off Natl Chasse & Faune Sauvage, Studies & Expertise DRE, F-01330 Birieux, France Univ Haifa, Dept Evolutionary & Environm Biol, IL-3498838 Haifa, Israel Western Ecosyst Technol Inc, Laramie, WY 82070 USA Aarhus Univ, Arctic Res Ctr, DK-8000 Aarhus C, Denmark Polish Acad Sci, Inst Nat Conservat, PL-31120 Krakow, Poland Univ Porto, CIBIO InBIO Associate Lab, P-4485661 Vairao, Portugal Univ Lisbon, Inst Super Agron, InBIO Associate Lab, Ctr Appl Ecol Prof Baeta Neves, P-1349017 Lisbon, Portugal Univ Lisbon, Fac Ciencias, Ctr Ecol Evolut & Environm Changes, P-1749016 Lisbon, Portugal Univ Calif Davis, Dept Environm Sci & Policy, Davis, CA 95616 USA Jackson Hole Conservat, Jackson, WY 83001 USA Univ Vet Med Hannover, Inst Terr & Aquat Wildlife Res, D-30173 Hannover, Germany Univ Calif Santa Cruz, Environm Studies Dept, Ctr Integrated Spatial Res, Santa Cruz, CA 95060 USA Univ Pretoria, Dept Zool & Entomol, ZA-0028 Hatfield, South Africa Colorado State Univ, Dept Fish Wildlife & Conservat Biol, Ft Collins, CO 80523 USA Tatra Natl Pk, PL-34500 Zakopane, Poland Sao Paulo State Univ, Anim Biol Postgra Program, BR-15054000 Sao Jose Do Rio Preto, SP, Brazil

Migratory behavior in ungulates has declined globally and understanding the causative factors (environmental change vs. human mediated) is needed to formulate effective management strategies. In the Jackson elk herd of northwest Wyoming, demographic differences between summer elk (Cervus elaphus) population segments have led to changes in migratory patterns over a 35-year time period. The proportion of short-distance migrants (SDM) has increased and the proportion of long-distance migrants (LDM) has concurrently declined. The probability of winter-captured elk on the National Elk Refuge being LDM decreased from 0.99 (95% CI = 0.97–1.00) to 0.59 (95% CI = 0.47–0.70) from 1978 to 2012. We tested 4 hypotheses that could contribute toward the decline in the LDM segment: behavioral switching from LDM to SDM, differential survival, harvest availability, and calf recruitment. Switching rates from LDM to SDM were very low (0.2% each elk-year). Survival rates were similar between LDM and SDM, although harvest availability was relatively low for SDM that tended to use areas close to human development during the hunting season. Average summer calf/cow ratios of LDM declined from 42 to 23 calves per 100 cows from 1978–1984 to 2006–2012. Further, during 2006–2012, LDM summer calf/cow ratios were less than half of SDM (23 vs. 47 calves per 100 cows). Our data suggest recruitment is the driving factor behind the declining proportion of LDM in this region. Effectiveness of altering harvest management strategies to conserve the LDM portion of the Jackson elk herd may be limited. Published 2015. This article is a U.S. Government work and is in the public domain in the USA.

As the extent and intensity of energy development in North America increases, so do disturbances to wildlife and the habitats they rely upon. Impacts to mule deer are of particular concern because some of the largest gas fields in the USA overlap critical winter ranges. Short-term studies of 2–3 years have shown that mule deer and other ungulates avoid energy infrastructure; however, there remains a common perception that ungulates habituate to energy development, and thus, the potential for a demographic effect is low. We used telemetry data from 187 individual deer across a 17-year period, including 2 years predevelopment and 15 years during development, to determine whether mule deer habituated to natural gas development and if their response to disturbance varied with winter severity. Concurrently, we measured abundance of mule deer to indirectly link behavior with demography. Mule deer consistently avoided energy infrastructure through the 15-year period of development and used habitats that were an average of 913 m further from well pads compared with predevelopment patterns of habitat use. Even during the last 3 years of study, when most wells were in production and reclamation efforts underway, mule deer remained >1 km away from well pads. The magnitude of avoidance behavior, however, was mediated by winter severity, where aversion to well pads decreased as winter severity increased. Mule deer abundance declined by 36% during the development period, despite aggressive onsite mitigation efforts (e.g. directional drilling and liquid gathering systems) and a 45% reduction in deer harvest. Our results indicate behavioral effects of energy development on mule deer are long term and may affect population abundance by displacing animals and thereby functionally reducing the amount of available habitat.

Partial migration occurs across a variety of taxa and has important ecological and evolutionary consequences. Among ungulates, studies of partially migratory populations have allowed researchers to compare and contrast performance metrics of migrants versus residents and examine how environmental factors influence the relative abundance of each. Such studies tend to characterize animals discretely as either migratory or resident, but we suggest that variable migration distances within migratory herds are an important and overlooked form of population structure, with potential consequences for animal fitness. We examined whether the variation in individual migration distances (20–264 km) within a single wintering population of mule deer (Odocoileus hemionus) was associated with several critical behavioral attributes of migration, including timing of migration, time allocation to seasonal ranges, and exposure to anthropogenic mortality risks. Both the timing of migration and the amount of time animals allocated to seasonal ranges varied with migration distance. Animals migrating long distances (150–250 km) initiated spring migration more than three weeks before than those migrating moderate (50–150 km) or short distances (

Resource selection functions (RSFs) are typically estimated by comparing covariates at a discrete set of "used" locations to those from an "available" set of locations. This RSF approach treats the response as binary and does not account for intensity of use among habitat units where locations were recorded. Advances in global positioning system (GPS) technology allow animal location data to be collected at fine spatiotemporal scales and have increased the size and correlation of data used in RSF analyses. We suggest that a more contemporary approach to analyzing such data is to model intensity of use, which can be estimated for one or more animals by relating the relative frequency of locations in a set of sampling units to the habitat characteristics of those units with count-based regression and, in particular, negative binomial (NB) regression. We demonstrate this NB RSF approach with location data collected from 10 GPS-collared Rocky Mountain elk (Cervus elaphus) in the Starkey Experimental Forest and Range enclosure. We discuss modeling assumptions and show how RSF estimation with NB regression can easily accommodate contemporary research needs, including: analysis of large GPS data sets, computational ease, accounting for among-animal variation, and interpretation of model covariates. We recommend the NB approach because of its conceptual and computational simplicity, and the fact that estimates of intensity of use are unbiased in the face of temporally correlated animal location data.

Summary 1. Well-informed management of harvested species requires understanding how changing ecological conditions affect demography and population dynamics, information that is lacking for many species. We have limited understanding of the relative influence of carnivores, harvest, weather and forage availability on elk Cervus elaphus demography, despite the ecological and economic importance of this species. We assessed adult female survival, a key vital rate for population dynamics, from 2746 radio-collared elk in 45 populations across western North America that experience wide variation in carnivore assemblage, harvest, weather and habitat conditions. 2. Proportional hazard analysis revealed that ‘baseline’ (i.e. not related to human factors) mortality was higher with very high winter precipitation, particularly in populations sympatric with wolves Canis lupus. Mortality may increase via nutritional stress and heightened vulnerability to predation in snowy winters. Baseline mortality was unrelated to puma Puma concolor presence, forest cover or summer forage productivity. 3. Cause-specific mortality analyses showed that wolves and all carnivore species combined had additive effects on baseline elk mortality, but only reduced survival by

Summary 1. Impermeable barriers to migration can greatly constrain the set of possible routes and ranges used by migrating animals. For ungulates, however, many forms of development are semi-permeable, and making informed management decisions about their potential impacts to the persistence of migration routes is difficult because our knowledge of how semi-permeable barriers affect migratory behaviour and function is limited. 2. Here, we propose a general framework to advance the understanding of barrier effects on ungulate migration by emphasizing the need to (i) quantify potential barriers in terms that allow behavioural thresholds to be considered, (ii) identify and measure behavioural responses to semi-permeable barriers and (iii) consider the functional attributes of the migratory landscape (e.g. stopovers) and how the benefits of migration might be reduced by behavioural changes. 3. We used global position system (GPS) data collected from two subpopulations of mule deer Odocoileus hemionus to evaluate how different levels of gas development influenced migratory behaviour, including movement rates and stopover use at the individual level, and intensity of use and width of migration route at the population level. We then characterized the functional landscape of migration routes as either stopover habitat or movement corridors and examined how the observed behavioural changes affected the functionality of the migration route in terms of stopover use. 4. We found migratory behaviour to vary with development intensity. Our results suggest that mule deer can migrate through moderate levels of development without any noticeable effects on migratory behaviour. However, in areas with more intensive development, animals often detoured from established routes, increased their rate of movement and reduced stopover use, while the overall use and width of migration routes decreased. 5. Synthesis and applications. In contrast to impermeable barriers that impede animal movement, semi-permeable barriers allow animals to maintain connectivity between their seasonal ranges. Our results identify the mechanisms (e.g. detouring, increased movement rates, reduced stopover use) by which semi-permeable barriers affect the functionality of ungulate migration routes and emphasize that the management of semi-permeable barriers may play a key role in the conservation of migratory ungulate populations.