Spatial ecology of the Vicuña (Lama vicugna) in a high Andean protected area.

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.

El estudio del uso del espacio en animales es fundamental para la conservación y gestión eficaz de sus poblaciones y hábitats silvestres en un mundo que cambia rápidamente, sin embargo muchas especies siguen estando mal descritas. Tal es el caso del estudio de ecología espacial de la vicuña, un camélido silvestre de tamaño mediano que tiene un papel crítico en la red trófica altoandina. Estudiamos el uso del espacio de 24 hembras adultas de vicuñas desde abril de 2014 hasta febrero de 2017 en el extremo sur del área de distribución de la especie. Las vicuñas mostraron una gran fidelidad en la ubicación de sus áreas de acción durante el período de estudio y compartieron gran parte de sus áreas de acción con otros grupos familiars. Las áreas de acción de las vicuñas en nuestro estudio fueron considerablemente más grandes que las estimaciones previas en todo el rango de la especie. Variacíon en factores ambientales y topográficos, y el riesgo asociado de depredación afectaron las distancias de las migraciones diarias de las vicuñas, pero no afectaron el tamaño del área de acción. Nuestro estudio reporta nuevos datos ecológicos sobre el uso del espacio de vicuñas que pueden informar los esfuerzos de conservación y manejo de esta especie y otros ungulados sociales.

Wherever I may roam-Human activity alters movements of red deer (Cervus elaphus) and elk (Cervus canadensis) across two continents.

Human activity and associated landscape modifications alter the movements of animals with consequences for populations and ecosystems worldwide. Species performing long-distance movements are thought to be particularly sensitive to human impact. Despite the increasing anthropogenic pressure, it remains challenging to understand and predict animals' responses to human activity. Here we address this knowledge gap using 1206 Global Positioning System movement trajectories of 815 individuals from 14 red deer (Cervus elaphus) and 14 elk (Cervus canadensis) populations spanning wide environmental gradients, namely the latitudinal range from the Alps to Scandinavia in Europe, and the Greater Yellowstone Ecosystem in North America. We measured individual-level movements relative to the environmental context, or movement expression, using the standardized metric Intensity of Use, reflecting both the directionality and extent of movements. We expected movement expression to be affected by resource (Normalized Difference Vegetation Index, NDVI) predictability and topography, but those factors to be superseded by human impact. Red deer and elk movement expression varied along a continuum, from highly segmented trajectories over relatively small areas (high intensity of use), to directed transitions through restricted corridors (low intensity of use). Human activity (Human Footprint Index, HFI) was the strongest driver of movement expression, with a steep increase in Intensity of Use as HFI increased, but only until a threshold was reached. After exceeding this level of impact, the Intensity of Use remained unchanged. These results indicate the overall sensitivity of Cervus movement expression to human activity and suggest a limitation of plastic responses under high human pressure, despite the species also occurring in human-dominated landscapes. Our work represents the first comparison of metric-based movement expression across widely distributed populations of a deer genus, contributing to the understanding and prediction of animals' responses to human activity.

Incorporating human dimensions is associated with better wildlife translocation outcomes.

Wildlife translocations are increasingly used to combat declining biodiversity worldwide. Successful translocation often hinges on coexistence between humans and wildlife, yet not all translocation efforts explicitly include human dimensions (e.g., economic incentives, education programs, and conflict reduction assistance). To evaluate the prevalence and associated outcomes of including human dimensions as objectives when planning translocations, we analyze 305 case studies from the IUCN's Global Re-Introduction Perspectives Series. We find that fewer than half of all projects included human dimension objectives (42%), but that projects including human dimension objectives were associated with improved wildlife population outcomes (i.e., higher probability of survival, reproduction, or population growth). Translocation efforts were more likely to include human dimension objectives if they involved mammals, species with a history of local human conflict, and local stakeholders. Our findings underscore the importance of incorporating objectives related to human dimensions in translocation planning efforts to improve conservation success.

Large carnivores avoid humans while prioritizing prey acquisition in anthropogenic areas.

Large carnivores are recovering in many landscapes where the human footprint is simultaneously growing. When carnivores encounter humans, the way they behave often changes, which may subsequently influence how they affect their prey. However, little research investigates the behavioural mechanisms underpinning carnivore response to humans. As a result, it is not clear how predator-prey interactions and their associated ecosystem processes will play out in the human-dominated areas into which carnivore populations are increasingly expanding. We hypothesized that humans would reduce predation risk for prey by disturbing carnivores or threatening their survival. Alternatively, or additionally, we hypothesized that humans would increase predation risk by providing forage resources that congregate herbivorous prey in predictable places and times. Using grey wolves Canis lupus in Jackson Hole, Wyoming, USA as a study species, we investigated 170 kill sites across a spectrum of human influences ranging from heavily restricted human activities on protected federal lands to largely unregulated activities on private lands. Then, we used conditional logistic regression to quantify how the probability of predation changed across varied types and amounts of human influences, while controlling for environmental characteristics and prey availability. Wolves primarily made kills in environmental terrain traps and where prey availability was high, but predation risk was significantly better explained with the inclusion of human influences than by environmental characteristics alone. Different human influences had different, and even converse, effects on the risk of wolf predation. For example, where prey were readily available, wolves preferentially killed animals far from motorized roads but close to unpaved trails. However, wolves responded less strongly to humans, if at all, where prey were scarce, suggesting they prioritized acquiring prey over avoiding human interactions. Overall, our work reveals that the effects of large carnivores on prey populations can vary considerably among different types of human influences, yet carnivores may not appreciably alter predatory behaviour in response to humans if prey are difficult to obtain. These results shed new light on the drivers of large carnivore behaviour in anthropogenic areas while improving understanding of predator-prey dynamics in and around the wildland-urban interface.

Fencing amplifies individual differences in movement with implications on survival for two migratory ungulates.

Fences have recently been recognized as one of the most prominent linear infrastructures on earth. As animals traverse fenced landscapes, they adjust movement behaviours 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 Behaviour Analysis, we classified individual movement responses upon encountering fences (i.e. barrier behaviours). We adopted the reaction norm framework to jointly quantify individual plasticity and behavioural types of barrier behaviours, as well as behaviour syndromes between barrier behaviours and animal space use. We also assessed whether barrier behaviours affect individual survival. Our results highlighted a high-level individual plasticity encompassing differences in the degree and direction of barrier behaviours 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 behaviours and space use and was negatively associated with individual survival. However, these relationships were not statistically significant for pronghorn. By integrating approaches from movement ecology and behavioural 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 behavioural shifts for wide-ranging animals in fenced landscapes.

Risk effects cascade up to an obligate scavenger.

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.

Dietary patterns of a versatile large carnivore, the puma (Puma concolor).

Large carnivores play critical roles in terrestrial ecosystems but have suffered dramatic range contractions over the past two centuries. Developing an accurate understanding of large carnivore diets is an important first step towards an improved understanding of their ecological roles and addressing the conservation challenges faced by these species.The puma is one of seven large felid species in the world and the only one native to the non-tropical regions of the New World. We conducted a meta-analysis of puma diets across the species' range in the Americas and assessed the impact of varying environmental conditions, niche roles, and human activity on puma diets. Pumas displayed remarkable dietary flexibility, consuming at least 232 different prey species, including one Critically Endangered and five Endangered species.Our meta-analysis found clear patterns in puma diets with changing habitat and environmental conditions. Pumas consumed more larger-bodied prey species with increasing distance from the equator, but consumption of medium-sized species showed the opposite trend.Puma diets varied with their realized niche; however, contrary to our expectations, puma consumption of large species did not change with their trophic position, and pumas consumed more small prey and birds as apex predators. Consumption of domestic species was negatively correlated with consumption of medium-sized wild species, a finding which underscores the importance of maintaining intact native prey assemblages.The tremendous dietary flexibility displayed by pumas represents both an opportunity and a challenge for understanding the puma's role in ecosystems and for the species' management and conservation. Future studies should explore the linkages between availability and selection of primary and other wild prey, and consequent impacts on predation of domestic species, in order to guide conservation actions and reduce conflict between pumas and people.

Carnivore Niche Partitioning in a Human Landscape.

AbstractTo 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 (Puma concolor), a dominant mesocarnivore (Lycalopex culpaeus), and a subordinate small carnivore (Lycalopex griseus)-in a human landscape featuring pastoralists and semidomestic carnivores (i.e., dogs, Canis familiaris). We found that dominant species had strong negative effects on the space use of subordinate ones, which ultimately created space for subordinate small carnivores. Humans and dogs were strictly diurnal, whereas the native carnivore community was nocturnal and exhibited high temporal overlap. Dietary overlap was high among the native carnivores, but dogs were trophically decoupled, largely because of human food subsidies. Our results show that in landscapes with evident human presence, temporal and dietary partitioning among native carnivores can be limited, leaving space as the most important axis to be partitioned among carnivores. We believe that these findings-the first to simultaneously assess all three critical niche axes among competing carnivores and humans and their associated species (i.e., domesticated carnivores)-are transferable to other carnivore communities in human-modified landscapes.

Cascading effects of a disease outbreak in a remote protected area.

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.

Limited sexual segregation in a dimorphic avian scavenger, the Andean condor.

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.

The plasticity of ungulate migration in a changing world.

Migratory ungulates are thought to be declining globally because their dependence on large landscapes renders them highly vulnerable to environmental change. Yet recent studies reveal that many ungulate species can adjust their migration propensity in response to changing environmental conditions to potentially improve population persistence. In addition to the question of whether to migrate, decisions of where and when to migrate appear equally fundamental to individual migration tactics, but these three dimensions of plasticity have rarely been explored together. Here, we expand the concept of migratory plasticity beyond individual switches in migration propensity to also include spatial and temporal adjustments to migration patterns. We develop a novel typological framework that delineates every potential change type within the three dimensions, then use this framework to guide a literature review. We discuss broad patterns in migratory plasticity, potential drivers of migration change, and research gaps in the current understanding of this trait. Our result reveals 127 migration change events in direct response to natural and human-induced environmental changes across 27 ungulate species. Species that appeared in multiple studies showed multiple types of change, with some exhibiting the full spectrum of migratory plasticity. This result highlights that multidimensional migratory plasticity is pervasive in ungulates, even as the manifestation of plasticity varies case by case. However, studies thus far have rarely been able to determine the fitness outcomes of different types of migration change, likely due to the scarcity of long-term individual-based demographic monitoring as well as measurements encompassing a full behavioral continuum and environmental gradient for any given species. Recognizing and documenting the full spectrum of migratory plasticity marks the first step for the field of migration ecology to employ quantitative methods, such as reaction norms, to predict migration change along environmental gradients. Closer monitoring for changes in migratory propensity, routes, and timing may improve the efficacy of conservation strategies and management actions in a rapidly changing world.

Drivers of site fidelity in ungulates.

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. 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. 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. 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. 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.

Where and when to hunt? Decomposing predation success of an ambush carnivore.

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-vicuña) 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.

Wave-like Patterns of Plant Phenology Determine Ungulate Movement Tactics.

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" [3-5]. 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.

Habitat complexity mediates the predator-prey space race.

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 vicuña 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 vicuñas 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-vicuña 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. Vicuña 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 vicuña habitat selection and utilization than in the canyon. Although pumas consistently selected for and utilized vegetative and topographic cover regardless of habitat distribution, vicuñas 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 vicuñas 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.

Integrating temporal refugia into landscapes of fear: prey exploit predator downtimes to forage in risky places.

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.

Plasticity in elk migration timing is a response to changing environmental conditions.

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.

Landscapes of Fear: Spatial Patterns of Risk Perception and Response.

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.

Migratory plasticity is not ubiquitous among large herbivores.

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.

Hidden cost of disease in a free-ranging ungulate: brucellosis reduces mid-winter pregnancy in elk.

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.