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The study of animal space use is fundamental to effective conservation and management of wildlife populations and habitats in a rapidly changing world, yet many species remain poorly described. Such is the case for the spatial ecology of the Vicuña–a medium-sized wild camelid that plays a critical role, both as a consumer and as prey, in the high Andean food web. We studied patterns of space use of 24 adult female vicuñas from April 2014 to February 2017 at the southern edge of its range. Vicuñas showed strong fidelity to their home range locations across the study period and shared large portions of their home ranges with vicuñas from other family groups. Vicuña home ranges in our study were considerably larger than previous estimates across the range of the species. Variation in environmental and terrain factors and the associated risk of predation affected vicuña diel migration distance but not home range size or overlap. Our study offers new ecological insights into vicuña space use that can inform conservation and management efforts of vicuñas and other social ungulates.

To minimize competitive overlap, carnivores modify one of their critical niche axes: space, time, or resources. However, we currently lack rules for how carnivore communities operate in human-dominated landscapes. We simultaneously quantified overlap in the critical niche axes of a simple carnivore community-an apex carnivore (

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.

Competition structures ecological communities. In carnivorans, competitive interactions are disproportionately costly to subordinate carnivores who must account for the risk of interspecific killing when foraging. Accordingly, missed opportunity costs for meso-carnivores imposed by risk can benefit the smallest-bodied competitors. However, the extent to which the risk perpetuates into spatial partitioning in hierarchically structured communities remains unknown. To determine how risk-avoidance behaviors shape the space-use of carnivore communities, we studied a simple community of carnivores in northern Patagonia, Argentina: pumas (Puma concolor; an apex carnivore), culpeo foxes (Lycalopex culpaeus; a meso-carnivore), and chilla foxes (Lycalopex griseus; a small carnivore). We used multi-species occupancy models to quantify the space use within the carnivore community and giving-up densities to understand the behaviors that structure space use. Notably, we applied an analytical framework that tests whether the actual or perceived risk of predation most strongly influences the space use of subordinate carnivores although accounting for their foraging and vigilance behaviors. We found that there was a dominance hierarchy from the apex carnivore through the meso-carnivore to the subordinate small carnivore, which was reflected in space. Although both meso- and small carnivores exhibited similar predator avoidance behavioral responses to apex carnivores, the habitat associations of apex carnivores only altered meso-carnivore space use. The biases in risk management we observed for meso-carnivores likely translates into stable co-existence of this community of competing carnivores. We believe our analytical framework can be extended to other communities to quantify the spatial-behavioral tradeoffs of risk.

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.

The effects of predation risk on prey populations have been studied extensively; yet, how risk is manifested in a trophically linked guild-scavengers-has been overlooked. Risk could be particularly consequential for obligate scavengers that are vulnerable while foraging and rely on carrion provisioned by, and shared with, apex predators. We investigated whether Andean condors (Vultur gryphus) respond to predation risk in a landscape where the main source of carrion are camelids killed by pumas (Puma concolor). We hypothesized that condors would exhibit different behavioral responses to predation risk while they search, encounter, and exploit carrion. We explored condor habitat selection while flying by tracking nine birds with satellite transmitters and monitored via camera traps 41 natural carcasses and 25 experimental carrion stations. We found that condors searched for carrion in areas with a high probability of occurrence of puma kills. However, condors avoided exploiting carrion in areas featuring tall vegetation and steep slopes-selected by pumas to stalk prey-suggesting that condors manage risk primarily through the identification of safe foraging sites prior to landing. Our finding that condors avoided foraging near stalking cover for pumas highlights the importance of risk effects beyond predator-prey interactions, particularly for obligate scavengers.

Disease outbreaks induced by humans increasingly threaten wildlife communities worldwide. Like predators, pathogens can be key top-down forces in ecosystems, initiating trophic cascades that may alter food webs. An outbreak of mange in a remote Andean protected area caused a dramatic population decline in a mammalian herbivore (the vicuña), creating conditions to test the cascading effects of disease on the ecological community. By comparing a suite of ecological measurements to pre-disease baseline records, we demonstrate that mange restructured tightly linked trophic interactions previously driven by a mammalian predator (the puma). Following the mange outbreak, scavenger (Andean condor) occurrence in the ecosystem declined sharply and plant biomass and cover increased dramatically in predation refuges where herbivory was historically concentrated. The evidence shows that a disease-induced trophic cascade, mediated by vicuña density, could supplant the predator-induced trophic cascade, mediated by vicuña behaviour, thereby transforming the Andean ecosystem.

Large carnivores like gray wolves (Canis lupus) play key roles in regulating ecosystem structure and function. After being functionally extirpated from the United States by the early 1900s, wolves have recently recolonized portions of their historic ranges and are increasingly coming into contact with a rapidly-growing human population. When carnivores encounter humans, the way they behave, and therefore the way they shape ecosystems, is likely to change. Unfortunately, our ability to predict how wolves will affect ecosystems in human-dominated areas is limited by an incomplete understanding of how and why carnivores respond to human influence. We are therefore investigating wolf kill sites across Jackson Hole, Wyoming, where we can disentangle the effects of multiple simultaneous human influences. Specifically, we are evaluating whether and how spatiotemporal patterns of wolf predation may change in response to unnatural physical infrastructure, disturbance from general human activity, potential threat of mortality, and human-altered prey distributions. Our ongoing field study will help managers anticipate effects of wolf predation in and around human-influenced areas while contributing novel information to theories of predation risk and predator-prey interactions. Featured photo by YNP on Flickr. https://flic.kr/p/HGfKqs

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.

The landscape of fear is an important driver of prey space use. However, prey can navigate the landscape of fear by exploiting temporal refuges from predation risk. We hypothesized that diel patterns of predator and prey movement and space use would be inversely correlated due to temporal constraints on predator habitat domain. Specifically, we evaluated habitat selection and activity of the vicuña and its only predator, the puma, during three diel periods: day, dawn/dusk, and night. Pumas selected the same habitats regardless of diel period-vegetated and rugged areas that feature stalking cover for pumas-but increased their activity levels during dawn/dusk and night when they benefit from reduced detection by prey. Vicuñas avoided areas selected by pumas and reduced activity at night, but selected vegetated areas and increased activity by day and dawn/dusk. Vicuña habitat selection and movement strategies appeared to reduce the risk of encountering pumas; movement rates of pumas and vicuñas were negatively correlated across the diel cycle, and habitat selection was negatively correlated during dawn/dusk and night. Our study shows that an ambush predator's temporal activity and space use patterns interact to create diel refugia and shape the antipredator behaviors of its prey. Importantly, it is likely the very nature of ambush predators' static habitat specificity that makes predator activity important to temporally varying perceptions of risk. Prey which depend on risky habitats for foraging appear to mitigate risk by feeding when they can more easily detect predators and when predators are least active.

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.

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

Sexual segregation is widely reported among sexually dimorphic species and generally attributed to intraspecific competition. Prey diversity and human activities can reinforce niche segregation by increasing resource heterogeneity. Here, we explored trophic and spatial sexual segregation in the only avian scavenger that exhibits pronounced sexual size dimorphism (up to 50% difference in body mass) and a highly despotic social system, the Andean condor (Vultur gryphus). We predicted that larger and dominant males would exclude smaller and subordinate females from high-quality resources, leading to sexual segregation particularly in human-dominated landscapes showing increased prey diversity. We compared resource use between females and males across six sites in Argentina featuring a range of prey diversity via stable isotopes analysis of molted feathers (n=141 individuals). We then focused on two sites featuring contrasting levels of prey diversity and quanti-fied assimilated diet via stable isotopes and space use via GPS monitoring (n=23 and 12 tagged individuals). We found no clear differences in isotopic niche space, individual variation in isotopic signature, or assimilated diet between females and males. However, there were differences in foraging locations between sexes, with females apparently using areas of fewer food resources more frequently than males. Local conditions defined the dynamics of fine-scale sexual differences in foraging sites; yet, unpredictable and ephemeral carrion resources likely prevent segregation by sexes at the landscape scale. Our study highlights complex dynamics of sexual segregation in vultures and the relevancy of analyses under multiple spatial–temporal scales to explore segregation in social species., This study was supported by Conservation Research and Education Opportunities International (CREOI), Hawk Mountain Sanctuary, the Rufford Foundation, the Department of Forest and Wildlife Ecology at the University of Wisconsin-Madison, The National Agency of Scientific and Technological Promotion of Argentina (PICT 0725-2014 and 2016-0354 BID), and Fundación BBVA (project BIOCON-08-126).

Spillover impacts pose challenges for the management of protected areas (PAs). The issue of external threats encroaching on PAs has long been recognized, but a corollary—that PA conservation can increase costs borne by neighboring governments or landowners—is less well appreciated. In some contexts, basic principles of fairness and cooperation suggest that PA users should help pay these costs. Several countries have developed mechanisms for distributing the costs of spillover impacts to PA users, but not the United States. Here, we investigate whether and how US park visitors could help address one type of spillover, the need for wildlife conservation efforts beyond park boundaries, using a case study of the Greater Yellowstone Ecosystem (GYE). We examine a “conservation fee” recently proposed in the Wyoming legislature, along with tax‐based alternatives. After exploring some costs of wildlife conservation in GYE, we estimate that a fee of up to $10 per vehicle could generate up to $13 million annually, and tax‐based approaches considerably more. We consider legal, political, and governance challenges, and ways to mitigate them. The GYE could serve as a demonstration site for visitor funding of cooperative, large‐landscape conservation, for potential future expansion in the US and beyond.

Sexual segregation is widely reported among sexually dimorphic species and generally attributed to intraspecific competition. Prey diversity and human activities can reinforce niche segregation by increasing resource heterogeneity. Here, we explored trophic and spatial sexual segregation in the only avian scavenger that exhibits pronounced sexual size dimorphism (up to 50% difference in body mass) and a highly despotic social system, the Andean condor (Vultur gryphus). We predicted that larger and dominant males would exclude smaller and subordinate females from high-quality resources, leading to sexual segregation particularly in human-dominated landscapes showing increased prey diversity. We compared resource use between females and males across six sites in Argentina featuring a range of prey diversity via stable isotopes analysis of molted feathers (n = 141 individuals). We then focused on two sites featuring contrasting levels of prey diversity and quantified assimilated diet via stable isotopes and space use via GPS monitoring (n = 23 and 12 tagged individuals). We found no clear differences in isotopic niche space, individual variation in isotopic signature, or assimilated diet between females and males. However, there were differences in foraging locations between sexes, with females apparently using areas of fewer food resources more frequently than males. Local conditions defined the dynamics of fine-scale sexual differences in foraging sites; yet, unpredictable and ephemeral carrion resources likely prevent segregation by sexes at the landscape scale. Our study highlights complex dynamics of sexual segregation in vultures and the relevancy of analyses under multiple spatial-temporal scales to explore segregation in social species.

Mobile species face an array of human threats across political boundaries, and their protection relies on identifying and prioritizing areas for conservation. Large avian scavengers are one of the widest ranging and most threatened species globally, and efforts to preserve them have come to the forefront of wildlife management. Vultures require access to functionally distinct habitats for roosting, foraging and flying, yet behavior-specific habitat modelling has been overlooked in management planning. Herein, we developed a spatial prioritization model for the threatened Andean condor (Vultur gryphus) that integrates activity-specific habitat selection across heterogeneous landscapes. We tracked 35 individuals in two regions of Argentina and Chile differing in topography and vegetation composition, and analyzed how landscape covariates influence where condors roost, forage and fly, while accounting for individual differences. We found that individuals responded differently to environmental covariates during each behavior, and identified regional differences for some covariates dependent on behavioral state. We also found important individual differences in habitat selection between birds inhabiting each region. We combined these results into an ensemble spatial prioritization model, and found that most areas of high priority for Andean condor conservation are not under protection. The strategic implementation of conservation measures in these priority areas could have important implications for the recovery of this species. Our study illustrates the value of integrating behavioral- and individual-specific habitat analyses into spatial conservation planning, and points to opportunities for effective management of threatened vultures., Financial support was provided by the Department of Forest and Wildlife Ecology - University of Wisconsin-Madison, Hawk Mountain Sanctuary, The Rufford Foundation, Mrs. Patricia Manigault, Agencia Nacional de Promoción Científica y Técnica (PICT-BID 0725/2014, Argentina), The Eppley Foundation for Research and Fundación BBVA (project BIOCON-08-126).

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.

Demonstrating disease impacts on the vital rates of free-ranging mammalian hosts typically requires intensive, long-term study. Evidence for chronic pathogens affecting reproduction but not survival is rare, but has the potential for wide-ranging effects. Accurately quantifying disease-associated reductions in fecundity is important for advancing theory, generating accurate predictive models, and achieving effective management. We investigated the impacts of brucellosis (Brucella abortus) on elk (Cervus canadensis) productivity using serological data from over 6,000 captures since 1990 in the Greater Yellowstone Ecosystem, USA. Over 1,000 of these records included known age and pregnancy status. Using Bayesian multilevel models, we estimated the age-specific pregnancy probabilities of exposed and naïve elk. We then used repeat-capture data to investigate the full effects of the disease on life history. Brucellosis exposure reduced pregnancy rates of elk captured in mid- and late-winter. In an average year, we found 60% of exposed 2-year-old elk were pregnant compared to 91% of their naïve counterparts (a 31 percentage point reduction, 89% HPDI=20%-42%), whereas exposed 3- to 9-year-olds were 7 percentage points less likely to be pregnant than naïve elk of their same age (89% HPDI=2%-11%). We found these reduced rates of pregnancy to be independent from disease-induced abortions, which afflict a portion of exposed elk. We estimate that the combination of reduced pregnancy by mid-winter and the abortions following mid-winter reduces the reproductive output of exposed female elk by 24%, which affects population dynamics to a similar extent as severe winters or droughts. Exposing hidden reproductive costs of disease is essential to avoid conflating them with the effects of climate and predation. Such reproductive costs cause complex population dynamics, and the magnitude of the effect we found should drive a strong selection gradient if there is heritable resistance.

Predator-prey games emerge when predators and prey dynamically respond to the behavior of one another, driving the outcomes of predator-prey interactions. Predation success is a function of the combined probabilities of encountering and capturing prey, which are influenced by both prey behavior and environmental features. While the relative importance of encounter and capture probabilities have been evaluated in a spatial framework, temporal variation in prey behavior and intrinsic catchability are likely to also affect the distribution of predation events. Using a single-predator-single-prey (puma-vicuna) system, we evaluated which factors predict predation events across both temporal and spatial dimensions of the components of predation by testing the prey-abundance hypothesis (predators select for high encounter probability) and the prey-catchability hypothesis (predators select for high relative capture probability) in time and space. We found that for both temporal and spatial analyses, neither the prey-abundance hypothesis nor the prey-catchability hypothesis alone predicted kill frequency or distribution; puma kill frequency was static throughout the diel cycle and pumas consistently selected a single habitat type when hunting, despite temporal and spatial variation in encounter rates and intrinsic catchability. Our integrated spatiotemporal analysis revealed that an interaction between time of day and habitat influences kill probability, suggesting that trade-offs in the temporal and spatial components of predation drive the probability of predation events. These findings reinforce the importance of examining both the temporal and spatial patterns of the components of predation, rather than unidimensional measures of predator or prey behavior, to comprehensively describe the feedbacks between predator and prey in the predator-prey game.

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.

Mitigation of large carnivore depredation is essential to increasing stakeholder support for human–carnivore coexistence. Lethal and non-lethal techniques are implemented by managers, livestock producers, and other stakeholders to reduce livestock depredations by large carnivores. However, information regarding the relative effectiveness of techniques commonly used to reduce livestock depredations is currently lacking. We evaluated 66 published, peer-reviewed research papers that quantitatively measured livestock depredation before and after employing 4 categories of lethal and non-lethal mitigation techniques (livestock husbandry, predator deterrents and removal, and indirect management of land or wild prey) to assess their relative effectiveness as livestock protection strategies. Effectiveness of each technique was measured as the reported percent change in livestock losses. Husbandry (42–100% effective) and deterrents (0–100% effective) demonstrated the greatest potential but also the widest variability in effectiveness in reducing livestock losses. Removal of large carnivores never achieved 100% effectiveness but exhibited the lowest variation (67–83%). Although explicit measures of effectiveness were not reported for indirect management, livestock depredations commonly decreased with sparser and greater distances from vegetation cover, at greater distances from protected areas, and in areas with greater wild prey abundance. Information on time duration of effects was available only for deterrents; a tradeoff existed between the effectiveness of tools and the length of time a tool remained effective. Our assessment revealed numerous sources of bias regarding the effectiveness of techniques as reported in the peer-reviewed literature, including a lack of replication across species and geographic regions, a focus on Canid carnivores in the United States, Europe, and Africa, and a publication bias toward studies reporting positive effects. Given these limitations, we encourage managers and conservationists to work with livestock producers to more consistently and quantitatively measure and report the impacts of mitigation techniques under a wider range of environmental, economic, and sociological conditions. © 2016 The Wildlife Society.

The ungulate–carnivore–vulture complex is a key trophic module of many terrestrial ecosystems, but one that is globally under threat. Few have explored cross-species dependencies in this module, and the degree to which vultures rely on trophic facilitation by apex carnivores is rarely known and almost never quantified. We investigated the importance of puma Puma concolor predation on its native camelid prey, vicuñas Vicugna vicugna and guanacos Lama guanicoe, in food provisioning for Andean condors Vultur gryphus in the high Andes of north-western Argentina. We evaluated the origin of wild food sources through carcass surveys. We quantified condor feeding habits via foraging observations and through the analysis of pellet contents and stable isotopes from moulted feathers. Of the 102 fresh camelid carcasses we monitored, nearly all (94%) resulted from puma predation, and the majority (85%) of camelid carcasses used by condors were killed by pumas. Camelids represented 88% of the prey items identified from 183 condor pellets, and isotopic analyses of moulted feathers from 86 individuals identified via multilocus genotyping revealed that camelids and Small livestock were the most important prey items, representing 45–58% and 28–38% of condor assimilated biomass, respectively. Synthesis and applications. Our results show that puma predation plays a key role in the foraging ecology of Andean condors, and highlight the importance of predatory processes that make carrion available to scavengers. We contend that targeting the conservation of ungulate–carnivore–vulture modules, rather than a species-specific approach, will be a more effective strategy to ensure the long-term persistence of Andean condors and other obligate scavengers. Fil: Perrig, Paula L.. University of Wisconsin; Estados Unidos Fil: Donadio, Emiliano. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Instituto de Investigaciones en Biodiversidad y Medioambiente. Universidad Nacional del Comahue. Centro Regional Universidad Bariloche. Instituto de Investigaciones en Biodiversidad y Medioambiente; Argentina Fil: Middleton, Arthur D.. University of Yale; Estados Unidos Fil: Pauli, Jonathan Nicholas. University of Wisconsin; Estados Unidos

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.

© 2019 Elsevier Ltd Animals experience varying levels of predation risk as they navigate heterogeneous landscapes, and behavioral responses to perceived risk can structure ecosystems. The concept of the landscape of fear has recently become central to describing this spatial variation in risk, perception, and response. We present a framework linking the landscape of fear, defined as spatial variation in prey perception of risk, to the underlying physical landscape and predation risk, and to resulting patterns of prey distribution and antipredator behavior. By disambiguating the mechanisms through which prey perceive risk and incorporate fear into decision making, we can better quantify the nonlinear relationship between risk and response and evaluate the relative importance of the landscape of fear across taxa and ecosystems.

The spatial relationship between predator and prey is often conceptualized as a behavioral response race, in which prey avoid predators while predators track prey. Limiting habitat types can create spatial anchors for prey or predators, influencing the likelihood that the predator or prey response will dominate. Joint spatial anchors emerge when predator and prey occupy similar feeding habitat domains and risk and reward become spatially conflated, confusing predictions of which player will win the space race. These spatial dynamics of risk-foraging trade-offs are often obscured by habitat heterogeneity and community complexity in large vertebrate systems, fueling ambiguity regarding the generality of predictions from predator-prey theory. To test how habitat distribution influences the predator-prey space race, we examine correlation in puma and vicuna habitat selection and space use at two sites, one of which generates a distinct risk-foraging trade-off at a joint spatial anchor. The distribution of vegetation, which serves as both forage for vicunas and stalking cover for pumas, differs between the sites; the llano contains a single central meadow that acts as a joint spatial anchor, while the canyon is characterized by more heterogeneous vegetation. Puma-vicuna habitat selection correlation was positive in the llano and negative in the canyon, and similarly, utilization distributions were more strongly correlated in the llano than the canyon. Vicuna locations occurred at higher values of puma habitat selection and utilization in the llano than in the canyon. Similarly, puma locations in the llano occurred at higher values of vicuna habitat selection and utilization than in the canyon. Although pumas consistently selected for and utilized vegetative and topographic cover regardless of habitat distribution, vicunas only selected against vegetation in the heterogeneous canyon site, reducing spatial correlation with pumas. Our work suggests a joint spatial anchor favors pumas in the space race due to the inability for vicunas to avoid crucial foraging habitat. The outcome of the predator-prey space race appears to be strongly informed by the distribution of habitat, whereby corresponding predictability of predator and prey favors predators in the spatial game.

Ramon & Cajal research contract from the Spanish Ministry of Economy, Industry and Competitiveness (grant number RYC-2015-18932), Van Eeden, Lily M., Eklund, Ann, Miller,Jennifer R. B., López Bao, José Vicente, Chapron, Guillaume, Cejtin, Mikael R., Crowther, Mathew S., Dickman, Christopher R., Frank, Jens, Krofe, Miha, Macdonald, David W., McManus, Jeannine, Meyer, Tara K., Middleton, Arthur D., Newsome, Thomas M., Ripple, William J., Ritchie, Euan G., Schmitz, Oswald J., Stoner, kelly J., Tourani, Mahdieh, Treves, Adrian

We quantified patterns of population genetic structure to help understand gene flow among elk populations across the Greater Yellowstone Ecosystem. We sequenced 596 base pairs of the mitochondrial control region of 380 elk from eight populations. Analysis revealed high mitochondrial DNA variation within populations, averaging 13.0 haplotypes with high mean gene diversity (0.85). The genetic differentiation among populations for mitochondrial DNA was relatively high (FST = 0.161; P = 0.001) compared to genetic differentiation for nuclear microsatellite data (FST = 0.002; P = 0.332), which suggested relatively low female gene flow among populations. The estimated ratio of male to female gene flow (mm/mf = 46) was among the highest we have seen reported for large mammals. Genetic distance (for mitochondrial DNA pairwise FST) was not significantly correlated with geographic (Euclidean) distance between

ARTHUR D. MIDDLETON, MATTHEW J. KAUFFMAN, DOUGLAS E. MCWHIRTER, JOHN G. COOK, RACHEL C. COOK, ABIGAIL A. NELSON, MICHAEL D. JIMENEZ, AND ROBERT W. KLAVER Wyoming Cooperative Fish and Wildlife Research Unit, Department of Zoology and Physiology, University of Wyoming, Laramie, Wyoming 82071 USA Program in Ecology, University of Wyoming, Laramie, Wyoming 82071 USA U.S. Geological Survey, Wyoming Cooperative Fish and Wildlife Research Unit, Department of Zoology and Physiology, University of Wyoming, Laramie, Wyoming 82071 USA Wyoming Game and Fish Department, Cody, Wyoming 82414 USA National Council for Air and Stream Improvement, Forestry and Range Sciences Laboratory, La Grande, Oregon 97850 USA U.S. Fish and Wildlife Service, Jackson, Wyoming 83001 USA U.S. Geological Survey, Iowa Cooperative Fish and Wildlife Research Unit and Department of Natural Resource Ecology and Management, Iowa State University, Ames, Iowa 50011 USA

Annual variation in winter severity and growing season vegetation dynamics appear to influence the demography of temperate herbivores but parsing winter from spring effects requires independent metrics of environmental conditions specific to each season. We tested for independence in annual variation amongst four common metrics used to describe winter severity and early growing season vegetation dynamics across the entire spatial distribution of elk (Cervus elaphus) in Wyoming from 1989 to 2006. Winter conditions and early growing season dynamics were correlated in a specific way. Winters with snow cover that ended early tended to be followed by early, but slow, rises in the normalized difference vegetation index (NDVI), while long winters with extended periods of snow cover were often followed by late and rapid rises in NDVI. Across the 35 elk ranges, 0.4–86.8 % of the variation in the rate of increase in NDVI’s in spring was explained by the date snow cover disappeared from SNOTEL stations. Because phenoclimatological metrics are correlated across seasons and shifting due to climate change, identifying environmental constraints on herbivore fitness, particularly migratory species, is more difficult than previously recognized.

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 (