Wolves,white‐tailed deer,and beaver: implications of seasonal prey switching for woodland caribou declines

Population increases of primary prey can negatively impact alternate prey populations via demographic and behavioural responses of a shared predator through apparent competition. Seasonal variation in prey selection patterns by predators also can affect secondary and incidental prey by reducing spatial separation. Global warming and landscape changes in Alberta's bitumen sands have resulted in prey enrichment, which is changing the large mammal predator–prey system and causing declines in woodland caribou Rangifer tarandus caribou populations. We assessed seasonal patterns of prey use and spatial selection by wolves Canis lupus in two woodland caribou ranges in northeastern Alberta, Canada, that have undergone prey enrichment following recent white‐tailed deer Odocoileus virginianus invasion. We determined whether risk of predation for caribou (incidental prey) and the proportion of wolf‐caused‐caribou mortalities varied with season. We found that wolves showed seasonal variation in primary prey use, with deer and beaver Castor canadensis being the most common prey items in wolf diet in winter and summer, respectively. These seasonal dietary patterns were reflected in seasonal wolf spatial resource selection and resulted in contrasting spatial relationships between wolves and caribou. During winter, wolf selection for areas used by deer maintained strong spatial separation between wolves and caribou, whereas wolf selection for areas used by beaver in summer increased the overlap with caribou. Changing patterns in wolf resource selection were reflected by caribou mortality patterns, with 76.2% of 42 adult female caribou mortalities occurring in summer. Understanding seasonal patterns of predation following prey enrichment in a multiprey system is essential when assessing the effect of predation on an incidental prey species. Our results support the conclusion that wolves are proximately responsible for woodland caribou population declines throughout much of their range.     

Ecologically based definition of seasons clarifies predator–prey interactions

Species interactions within food webs are driven by multiple constraints, including those imposed by seasonal changes in the environment. Ecologically sound definitions of seasons may therefore be a prerequisite for clarifying predator prey interactions. Most studies define biological seasons based on fixed schedules or on temporal changes in a single movement measurement. We used a novel clustering approach based on homogeneous space‐use patterns of GPS‐collared animals to reveal 7 biological seasons for caribou Rangifer tarandus caribou, and 5 for both moose Alces alces and grey wolves Canis lupus interacting in a boreal ecosystem. Subsequent evaluation of niche overlap showed that, as predicted, wolves had a stronger spatio‐temporal connection with moose, its main prey, than with caribou. Movement constraints and limiting resource distributions similarly affected all species in some instances, but also caused temporal changes in the extent of niche overlap between wolves and its two prey. The risk that caribou faced was not only linked to the niche overlap with wolves, but also to the extent of wolf‐moose niche overlap during the same period. Food‐web properties emerged from the analysis, with temporal changes in relative niche overlap reflecting the strength of trophic interactions during the year. Our study demonstrates how the study of trophic interactions can benefit from comprehensive definitions of biological seasons.     

Invading white-tailed deer change wolf–caribou dynamics in northeastern Alberta

Human-caused habitat change has been implicated in current woodland caribou (Rangifer tarandus caribou) population declines across North America. Increased early seral habitat associated with industrial footprint can result in an increase in ungulate densities and subsequently those of their predator, wolves (Canis lupus). Higher wolf densities can result in increased encounters between wolves and caribou and consequently higher caribou mortality. We contrasted changes in moose (Alces alces) and deer (Odocoileus spp.) densities and assessed their effects on wolf–caribou dynamics in northeastern Alberta, Canada, pre (1994–1997) versus post (2005–2009) major industrial expansion in the region. Observable white-tailed deer (O. virginianus) increased 17.5-fold but moose remained unchanged. Wolf numbers also increased from approximately 6–11.5/1,000 km². Coincident with these changes, spatial overlap between wolf pack territories and caribou range was high relative to the mid-1990s. The high number of wolf locations in caribou range suggests that forays were not merely exploratory, but rather represented hunting forays and denning locations. Scat analysis indicated that wolf consumption of moose declined substantively during this time period, whereas use of deer increased markedly and deer replaced moose as the primary prey of wolves. Caribou increased 10-fold in the diet of wolves and caribou population trends in the region changed from stable to declining. Wolf use of beaver (Castor canadensis) increased since the mid-1990s. We suggest that recent declines in woodland caribou populations in the southerly extent of their range have occurred because high deer densities resulted in a numeric response by wolves and consequently higher incidental predation on caribou. Our results indicate that management actions to conserve caribou must now include deer in primary prey and wolf reduction programs. © 2010 The Wildlife Society     

Trophic consequences of terrestrial eutrophication for a threatened ungulate

Changes in primary productivity have the potential to substantially alter food webs, with positive outcomes for some species and negative outcomes for others. Understanding the environmental context and species traits that give rise to these divergent outcomes is a major challenge to the generality of both theoretical and applied ecology. In aquatic systems, nutrient-mediated eutrophication has led to major declines in species diversity, motivating us to seek terrestrial analogues using a large-mammal system across 598 000 km² of the Canadian boreal forest. These forests are undergoing some of the most rapid rates of land-use change on Earth and are home to declining caribou (Rangifer tarandus caribou) populations. Using satellite-derived estimates of primary productivity, coupled with estimates of moose (Alces alces) and wolf (Canis lupus) abundance, we used path analyses to discriminate among hypotheses explaining how habitat alteration can affect caribou population growth. Hypotheses included food limitation, resource dominance by moose over caribou, and apparent competition with predators shared between moose and caribou. Results support apparent competition and yield estimates of wolf densities (1.8 individuals 1000 km⁻²) above which caribou populations decline. Our multi-trophic analysis provides insight into the cascading effects of habitat alteration from forest cutting that destabilize terrestrial predator–prey dynamics. Finally, the path analysis highlights why conservation actions directed at the proximate cause of caribou decline have been more successful in the near term than those directed further along the trophic chain.     

Unveiling trade‐offs in resource selection of migratory caribou using a mechanistic movement model of availability

Habitat selection is a multi‐level, hierarchical process that should be a key component in the balance between food acquisition and predation risk avoidance (food–predation trade‐off). However, to date, studies have not fully elucidated how fine‐ and broad‐scale habitat decisions by individual prey can help balance food versus risk. We studied broad‐scale habitat selection by Newfoundland caribou Rangifer tarandus, focusing on trade‐offs between predation risk versus access to forage during the calving and post‐calving period. We improved traditional measures of habitat availability by incorporating fine‐scale movement patterns of caribou into the availability kernel, thus enabling separation of broad and fine scales of selection. Remote sensing and field surveys served to create a spatio‐temporal model of forage availability, whereas GPS telemetry locations from 66 black bears Ursus americanus and 59 coyotes Canis latrans provided models of predation risk. We then used GPS telemetry locations from 114 female caribou to assess food–predation trade‐offs through the prism of our refined model of caribou habitat availability. We noted that migratory movements of caribou were oriented mainly towards habitats with abundant forage and lower risk of bear and (to a lesser extent) coyote encounter. These findings were generally consistent across caribou herds and would not have been evident had we used traditional methods instead of our refined model when estimating habitat availability. We interpret these findings in the context of stereotypical migratory behaviour observed in Newfoundland caribou, which occurs despite the extirpation of wolves Canis lupus nearly a century ago. We submit that caribou are able to balance food acquisition against predation risk using a complex set of factors involving both finer and broader scale selection. Accordingly, our study provides a strong argument for using refined habitat availability estimates when assessing food–predation trade‐offs.     

Coyote Habitat Selection and Management Implications for the Gaspésie Caribou

ABSTRACT Anthropogenic disturbances can promote establishment and growth of predator populations in areas where secondary prey can then become threatened. In this study, we investigated habitat selection of eastern coyotes (Canis latrans), a relatively new predator in the vicinity of an endangered population of caribou (Rangifer tarandus caribou). We hypothesized that coyotes in the boreal forest depend mainly on disturbed habitat, particularly that of anthropogenic origin, because these habitats provide increased food accessibility. Coyotes would likely take advantage of moose (Alces alces) carcasses, berries, and snowshoe hares (Lepus americanus) found in open habitats created by logging. To test these predictions, we described coyote diet and habitat selection at different spatial and temporal levels and then compared resource availability between habitats. To do so, we installed Global Positioning System radiocollars on 23 individual coyotes in the Gaspésie Peninsula, eastern Québec, Canada. Coyotes selected clear-cuts of 5–20 years and avoided mature coniferous forests both at the landscape and home-range levels. Clear-cuts of 5–20 years were found to contain a high availability of moose carcasses and berries, and vulnerability of snowshoe hares is known to increase in clear-cuts. The importance of these 3 food resources was confirmed by the characteristics of core areas used by coyotes and diet analysis. Moose remains were found at 45% of core areas and coyote diet comprised 51% moose on an annual basis. Anthropogenic disturbances in the boreal forest thus seem to benefit coyotes. Our results indicated that the relationship between coyotes and caribou likely involves spillover predation. This knowledge allows managers to consider spillover predation by coyotes as a possible threat for endangered caribou population when the predator depends mainly on habitat of anthropogenic origin and to suggest methods to alleviate it when developing management plans.     

Multiple spatial-scale resource selection function models in relation to human disturbance for moose in northeastern China

The moose (Alces alces cameloides) population in northeastern China is on the southernmost edge of its distribution in Asia. A survey was conducted to determine moose resource selection and the effects of human disturbance on moose in a study area of 20,661 ha located on the northwestern slope of the Lesser Khingan Mountains, located in northeastern China. Predictive models of resource selection were developed using logistic and autologistic regression. All models considered resource variable selection at two spatial scales, patch and landscape. At the patch scale, moose preferred larger birch (Betula platyphylla) patches, but avoided larger tamarack (Larix gmelinii) patches. At the landscape scale, moose preferred higher densities of tamarack patches, i.e., heterogeneity of tamarack stands, selected areas with more abundant annual shoots, terrain conducive to better concealment, higher altitudes and areas saturated with soil moisture. Roads and forest harvest intervals were identified as important human disturbance factors. This is the first time that moose have been reported to avoid roads, and the avoidance distance was nearly 3 km. We believe that in this region moose under the influence of roads are behaviorally plastic, compared with the indifference of moose to the presence of roads in other regions. Moose avoided forest areas logged more than 3 years previously and preferred areas logged 1–2 years previously. In addition, it may be necessary to monitor the effect of the dynamic of density of roe deer on the spatial distribution of the moose population.     

Understanding predation risk and individual variation in risk avoidance for threatened boreal caribou

Predation risk is a driver of species’ distributions. Animals can increase risk avoidance in response to fluctuations in predation risk, but questions remain regarding individual variability and the capacity to respond to changes in spatial risk across human‐altered landscapes. In northeast British Columbia, Canada, boreal caribou populations declined as roads and seismic lines have increased, which are theorized to increase gray wolf predation. Our goal was to model risk and to evaluate individual variability and the development of risk perception by examining individual risk avoidance in response to reproductive status and age. We used locations from collared caribou and wolves to identify landscape features associated with the risk of a potential wolf‐caribou encounter and risk of being killed given an encounter. We built resource selection functions to estimate individual responses to risk. We used general linear regressions to evaluate individual risk and linear feature avoidance as a function of age and reproductive status (calf or no calf). Linear features increased the risk of encounter. Older caribou and caribou with calves demonstrated stronger avoidance of the risk of encounter and roads, but weaker avoidance in late summer to the risk of being killed relative to younger and calf‐less individuals. Mechanisms explaining the inverse relationships between the risk of encounter and risk of being killed are uncertain, but it is conceivable that caribou learn to avoid the risk of encounter and roads. Responses by females with vulnerable calves to the risk of encounter and risk of being killed might be explained by a trade‐off between these two risk types and a prioritization on the risk of encounter. Despite the capacity to alter their responses to risk, the global decline in Rangifer populations (caribou and wild reindeer) suggests these behaviors are insufficient to mitigate the impacts of anthropogenic disturbances.     

Trophic niche partitioning between two prey and their incidental predators revealed various threats for an endangered species

Documenting trophic niche partitioning and resource use within a community is critical to evaluate underlying mechanisms of coexistence, competition, or predation. Detailed knowledge about foraging is essential as it may influence the vital rates, which, in turn, can affect trophic relationships between species, and population dynamics. The aims of this study were to evaluate resource and trophic niche partitioning in summer/autumn between the endangered Atlantic‐Gaspésie caribou (Rangifer tarandus caribou) population, moose (Alces americanus) and their incidental predators, the black bear (Ursus americanus) and coyote (Canis latrans), and to quantify the extent to which these predators consumed caribou. Bayesian isotopic analysis showed a small overlap in trophic niche for the two sympatric ungulates suggesting a low potential for resource competition. Our results also revealed that caribou occupied a larger isotopic niche area than moose, suggesting a greater diversity of resources used by caribou. Not surprisingly, coyotes consumed mainly deer (Odocoileus virginianus), moose, snowshoe hare (Lepus americanus), and occasionally caribou, while bears consumed mainly vegetation and, to a lesser extent, moose and caribou. As coyotes and bears also feed on plant species, we documented trophic niche overlap between caribou and their predators, as searching for similar resources can force them to use the same habitats and thus increase the encounter rate and, ultimately, mortality risk for caribou. Although the decline in the Gaspésie caribou population is mostly driven by habitat‐mediated predation, we found evidence that the low level of resource competition with moose, added to the shared resources with incidental predators, mainly bears, may contribute to jeopardize the recovery of this endangered caribou population. Highlighting the trophic interaction between species is needed to establish efficient conservation and management strategies to insure the persistence of endangered populations. The comparison of trophic niches of species sharing the same habitat or resources is fundamental to evaluate the mechanisms of coexistence or competition and eventually predict the consequences of ecosystem changes in the community.     

Behavioral “bycatch” from camera trap surveys yields insights on prey responses to human‐mediated predation risk

Human disturbance directly affects animal populations and communities, but indirect effects of disturbance on species behaviors are less well understood. For instance, disturbance may alter predator activity and cause knock‐on effects to predator‐sensitive foraging in prey. Camera traps provide an emerging opportunity to investigate such disturbance‐mediated impacts to animal behaviors across multiple scales. We used camera trap data to test predictions about predator‐sensitive behavior in three ungulate species (caribou Rangifer tarandus; white‐tailed deer, Odocoileus virginianus; moose, Alces alces) across two western boreal forest landscapes varying in disturbance. We quantified behavior as the number of camera trap photos per detection event and tested its relationship to inferred human‐mediated predation risk between a landscape with greater industrial disturbance and predator activity and a “control” landscape with lower human and predator activity. We also assessed the finer‐scale influence on behavior of variation in predation risk (relative to habitat variation) across camera sites within the more disturbed landscape. We predicted that animals in areas with greater predation risk (e.g., more wolf activity, less cover) would travel faster past cameras and generate fewer photos per detection event, while animals in areas with less predation risk would linger (rest, forage, investigate), generating more photos per event. Our predictions were supported at the landscape‐level, as caribou and moose had more photos per event in the control landscape where disturbance‐mediated predation risk was lower. At a finer‐scale within the disturbed landscape, no prey species showed a significant behavioral response to wolf activity, but the number of photos per event decreased for white‐tailed deer with increasing line of sight (m) along seismic lines (i.e., decreasing visual cover), consistent with a predator‐sensitive response. The presence of juveniles was associated with shorter behavioral events for caribou and moose, suggesting greater predator sensitivity for females with calves. Only moose demonstrated a positive behavioral association (i.e., longer events) with vegetation productivity (16‐day NDVI), suggesting that for other species bottom‐up influences of forage availability were generally weaker than top‐down influences from predation risk. Behavioral insights can be gleaned from camera trap surveys and provide complementary information about animal responses to predation risk, and thus about the indirect impacts of human disturbances on predator–prey interactions.     

Multi-Scale Foraging Habitat Use and Interactions by Sympatric Cervids in Northeastern China

Abstract: Moose (Alces alces) and roe deer (Capreolus pygargus) are sympatric in the forest region of northeastern China. Using univariate analyses of feeding sign data, we found the 2 species were positively associated, but there were distinctions between their use of forage resources across landscape, patch, and microhabitat scales. We used resource selection function models to predict the influence of environmental covariates on moose and roe deer foraging; we detected covariate effects at the landscape and microhabitat scales but not at the patch scale. Forage resources used by the 2 species were similar, but moose used wetter areas and more low-visibility habitats than did roe deer, which strongly avoided areas with sparse vegetation. Both species were influenced by forage abundance and distribution at the microhabitat scale but exhibited differences in intensity of use of plant species and microhabitats. Moose used areas with deeper snow and avoided hiding cover; roe deer avoided areas with higher total basal areas of tree stems and preferred areas with high plant species richness. For moose, there was a trade-off in the use of concealment cover between the landscape and microhabitat scales. We detected avoidance by moose of roads where roe deer occurred. Roe deer exhibited more capacity for coping with human disturbance and interspecific interaction. In areas similar to our study area, road closures and suppression of roe deer near roads within 3–5 years postlogging may benefit moose. Furthermore, a mosaic of areas with different logging intervals may contribute to spatial separation of moose and roe deer and promote their coexistence.     

The influence of landscape matrix on isolated patch use by wide‐ranging animals: conservation lessons for woodland caribou

For conservation purposes, it is important to design studies that explicitly quantify responses of focal species to different land management scenarios. Here, we propose an approach that combines the influence of landscape matrices with the intrinsic attributes of remaining habitat patches on the space use behavior of woodland caribou (Rangifer tarandus caribou), a threatened subspecies of Rangifer. We sought to link characteristics of forest remnants and their surrounding environment to caribou use (i.e., occurrence and intensity). We tracked 51 females using GPS telemetry north of the Saguenay River (Québec, Canada) between 2004 and 2010 and documented their use of mature forest remnants ranging between 30 and ~170 000 ha in a highly managed landscape. Habitat proportion and anthropogenic feature density within incremental buffer zones (from 100 to 7500 m), together with intrinsic residual forest patch characteristics, were linked to caribou GPS location occurrence and density to establish the range of influence of the surrounding matrix. We found that patch size and composition influence caribou occurrence and intensity of use within a patch. Patch size had to reach approximately 270 km² to attain 75% probability of use by caribou. We found that small patches (<100 km²) induced concentration of caribou activities that were shown to make them more vulnerable to predation and to act as ecological traps. Woodland caribou clearly need large residual forest patches, embedded in a relatively undisturbed matrix, to achieve low densities as an antipredator strategy. Our patch‐based methodological approach, using GPS telemetry data, offers a new perspective of space use behavior of wide‐ranging species inhabiting fragmented landscapes and allows us to highlight the impacts of large scale management. Furthermore, our study provides insights that might have important implications for effective caribou conservation and forest management.     

Foraging strategies by omnivores: are black bears actively searching for ungulate neonates or are they simply opportunistic predators?

Omnivores feed on animals with dynamic distributions and on plants with static distributions. The search tactics they adopt will not only define the risk for the targeted prey, but also for other prey that may be consumed when encountered. The potential impact of omnivores on the dynamics of multi‐prey systems thus depends on resource selection and on the tactics used to find their prey. We present an approach that can clarify the foraging decisions of omnivores by combining analyses of habitat selection, local residency time, and interpatch movements. We use this framework to evaluate whether predation by omnivorous black bears on ungulate neonates resulted from an active search or from incidental encounters. We monitored 12 bears, 22 forest‐dwelling caribou, and 36 moose during calving seasons. We estimated the spatial patterns in relative occurrence probability of ungulate neonates using Resource Selection Functions (RSFs). We also mapped plant abundance from vegetation surveys. RSF were then built to assess the link between bear distribution and the distribution of these three food types (vegetation, moose calves, caribou fawns). We further evaluated the search tactic used by bears that led to this spatial dependency by exploring patterns of residency times and interpatch movements. Bears did not select areas with a high probability of encounter with neonates, but selected areas with abundant vegetation. Surprisingly, bears displayed shorter residency times in vegetation‐rich areas. The selection for vegetation‐rich areas was therefore achieved by moving preferentially, but frequently, between areas offering abundant vegetation. Such frequent interpatch movements could result in high rates of fortuitous encounters with neonates, even if bears are not actively searching for them. To mitigate the impacts of forest harvesting on threatened caribou populations, vegetation‐rich areas selected by bears (e.g. roadsides) should be segregated from large patches of mature conifer forest suitable for caribou.     

Modeling Cumulative Effects of Climate and Development on Moose,Wolf, and Caribou Populations

Wildlife models focused solely on a single strong influence (e.g., habitat components, wildlife harvest) are limited in their ability to detect key mechanisms influencing population change. Instead, we propose integrated modeling in the context of cumulative effects assessment using multispecies population dynamics models linked to landscape-climate simulation at large spatial and temporal scales. We developed an integrated landscape and population simulation model using ALCES Online as the model-building platform, and the model accounted for key ecological components and relationships among moose (Alces alces), grey wolves (Canis lupus nubilus), and woodland caribou (Rangifer tarandus caribou) in northern Ontario, Canada. We simulated multiple scenarios over 5 decades (beginning 2020) to explore sensitivity to climate change and land use and assessed effects at multiple scales. The magnitude of effect and the relative importance of key factors (climate change, roads, and habitat) differed depending on the scale of assessment. Across the full extent of the study area (654,311km² [ecozonal scale]), the caribou population declined by 26% largely because of climate change and associated predator-prey response, which led to caribou range recession in the southern part of the study area. At the caribou range scale (108,378 km²), which focused on 2 herds in the northern part of the study area, climate change led to a 10% decline in the population and development led to an additional 7% decline. At the project scale (8,331 km²), which was focused more narrowly on the landscape surrounding 4 proposed mines, the caribou population declined by 29% largely in response to simulated development. Given that observed caribou population dynamics were sensitive to the cumulative effects of climate change, land use, interspecific interactions, and scale, insights from the analysis might not emerge under a less complex model. Our integrated modeling framework provides valuable support for broader regional assessments, including estimation of risk to caribou and Indigenous food security, and for developing and evaluating potential caribou recovery strategies. © 2021 The Authors. The Journal of Wildlife Management published by Wiley Periodicals LLC on behalf of The Wildlife Society.     

End of the road: Short-term responses of a large mammal community to forest road decommissioning

Anthropogenic disturbances are increasing worldwide, causing wildlife habitat loss, alteration, and fragmentation. In Canada, the decommissioning of linear anthropogenic structures is identified as a promising tool to restore the habitat of threatened populations of boreal caribou (Rangifer tarandus caribou) by reducing food availability for alternate prey and decreasing encounter probabilities with predators. In this study, we monitored the use of 40 km of decommissioned forest roads by caribou, gray wolves (Canis lupus), black bears (Ursus americanus), and moose (Alces americanus) 1–3 years after reclamation, using 232 motion-activated camera traps. We compared four additive treatments (meaning that each successive treatment included the treatment prior): closing the road to human access, decompacting its soil, planting black spruce (Picea mariana) trees, and adding enriched soil. We assessed the influence of treatments, use by other large mammals, and characteristics of the surrounding environment on road use by the four species. Caribou used the planted treatment (which also included closing and decompacting) more than the closed-only (reference) treatment, but treatments did not influence the use of decommissioned roads by bears and moose. We could not assess the use of treated roads by wolves because of low sample size. Road use by caribou declined with local moose density, but increased with local bear density. Caribou were observed more frequently on roads surrounded by regenerating and mature coniferous stands; caribou also preferentially used roads surrounded by wetlands. Our results suggest that the treatment combining road closure, soil decompaction, and tree planting could be beneficial to caribou, highlighting the relevance of including active restoration efforts in caribou conservation programs. We recommend that such a treatment be added to road decommissioning protocols for the conservation of caribou, alongside broad-scale habitat protection.     

The landscape of fear as an emergent property of heterogeneity: Contrasting patterns of predation risk in grassland ecosystems

The likelihood of encountering a predator influences prey behavior and spatial distribution such that non‐consumptive effects can outweigh the influence of direct predation. Prey species are thought to filter information on perceived predator encounter rates in physical landscapes into a landscape of fear defined by spatially explicit heterogeneity in predation risk. The presence of multiple predators using different hunting strategies further complicates navigation through a landscape of fear and potentially exposes prey to greater risk of predation. The juxtaposition of land cover types likely influences overlap in occurrence of different predators, suggesting that attributes of a landscape of fear result from complexity in the physical landscape. Woody encroachment in grasslands furnishes an example of increasing complexity with the potential to influence predator distributions. We examined the role of vegetation structure on the distribution of two avian predators, Red‐tailed Hawk (Buteo jamaicensis) and Northern Harrier (Circus cyaneus), and the vulnerability of a frequent prey species of those predators, Northern Bobwhite (Colinus virginianus). We mapped occurrences of the raptors and kill locations of Northern Bobwhite to examine spatial vulnerability patterns in relation to landscape complexity. We use an offset model to examine spatially explicit habitat use patterns of these predators in the Southern Great Plains of the United States, and monitored vulnerability patterns of their prey species based on kill locations collected during radio telemetry monitoring. Both predator density and predation‐specific mortality of Northern Bobwhite increased with vegetation complexity generated by fine‐scale interspersion of grassland and woodland. Predation pressure was lower in more homogeneous landscapes where overlap of the two predators was less frequent. Predator overlap created areas of high risk for Northern Bobwhite amounting to 32% of the land area where landscape complexity was high and 7% where complexity was lower. Our study emphasizes the need to evaluate the role of landscape structure on predation dynamics and reveals another threat from woody encroachment in grasslands.     

Effects of human‐induced prey depletion on large carnivores in protected areas: Lessons from modeling tiger populations in stylized spatial scenarios

Prey depletion is a major threat to the conservation of large carnivore species globally. However, at the policy‐relevant scale of protected areas, we know little about how the spatial distribution of prey depletion affects carnivore space use and population persistence. We developed a spatially explicit, agent‐based model to investigate the effects of different human‐induced prey depletion experiments on the globally endangered tiger (Panthera tigris) in isolated protected areas—a situation that prevails throughout the tiger's range. Specifically, we generated 120 experiments that varied the spatial extent and intensity of prey depletion across a stylized (circle) landscape (1,000 km²) and Nepal's Chitwan National Park (~1,239 km²). Experiments that created more spatially homogenous prey distributions (i.e., less prey removed per cell but over larger areas) resulted in larger tiger territories and smaller population sizes over time. Counterintuitively, we found that depleting prey along the edge of Chitwan National Park, while decreasing tiger numbers overall, also decreased female competition for those areas, leading to lower rates of female starvation. Overall our results suggest that subtle differences in the spatial distributions of prey densities created by various human activities, such as natural resource‐use patterns, urban growth and infrastructure development, or conservation spatial zoning might have unintended, detrimental effects on carnivore populations. Our model is a useful planning tool as it incorporates information on animal behavioral ecology, resource spatial distribution, and the drivers of change to those resources, such as human activities.     

Space use and social association in a gregarious ungulate: Testing the conspecific attraction and resource dispersion hypotheses

Animals use a variety of proximate cues to assess habitat quality when resources vary spatiotemporally. Two nonmutually exclusive strategies to assess habitat quality involve either direct assessment of landscape features or observation of social cues from conspecifics as a form of information transfer about forage resources. The conspecific attraction hypothesis proposes that individual space use is dependent on the distribution of conspecifics rather than the location of resource patches, whereas the resource dispersion hypothesis proposes that individual space use and social association are driven by the abundance and distribution of resources. We tested the conspecific attraction and the resource dispersion hypotheses as two nonmutually exclusive hypotheses explaining social association and of adult female caribou (Rangifer tarandus). We used location data from GPS collars to estimate interannual site fidelity and networks representing home range overlap and social associations among individual caribou. We found that home range overlap and social associations were correlated with resource distribution in summer and conspecific attraction in winter. In summer, when resources were distributed relatively homogeneously, interannual site fidelity was high and home range overlap and social associations were low. Conversely, in winter when resources were distributed relatively heterogeneously, interannual site fidelity was low and home range overlap and social associations were high. As access to resources changes across seasons, caribou appear to alter social behavior and space use. In summer, caribou may use cues associated with the distribution of forage, and in winter caribou may use cues from conspecifics to access forage. Our results have broad implications for our understanding of caribou socioecology, suggesting that caribou use season‐specific strategies to locate forage. Caribou populations continue to decline globally, and our finding that conspecific attraction is likely related to access to forage suggests that further fragmentation of caribou habitat could limit social association among caribou, particularly in winter when access to resources may be limited.     

Heading for the hills? Evaluating spatial distribution of woodland caribou in response to a growing anthropogenic disturbance footprint

Anthropogenic landscape change (i.e., disturbance) is recognized as an important factor in the decline and extirpation of wildlife populations. Understanding and monitoring the relationship between wildlife distribution and disturbance is necessary for effective conservation planning. Many studies consider disturbance as a covariate explaining wildlife behavior. However, we propose that there are several advantages to considering the spatial relationship between disturbance and wildlife directly using utilization distributions (UDs), including objective assessment of the spatially explicit overlap between wildlife and disturbance, and the ability to track trends in this relationship over time. Here, we examined how central mountain woodland caribou (Rangifer tarandus caribou) distribution changed over time in relation to (i) anthropogenic disturbance, baseline range (defined using telemetry data from 1998 to 2005), and alpine habitat; and (ii) interannual climate variation (North Pacific Index; NPI). We developed seasonal UDs for caribou in west‐central Alberta and east‐central British Columbia, Canada, monitored with GPS collars between 1998 and 2013. We mapped the cumulative annual density of disturbance features within caribou range and used indices of overlap to determine the spatial relationship and trend between caribou UDs, anthropogenic disturbance, baseline range, alpine habitat, and the NPI. Anthropogenic disturbance increased over time, but the overlap between caribou UDs and disturbance did not. Caribou use of alpine habitat during spring, fall, and late winter increased over time, concurrent with a decrease in use of baseline range. Overlap between caribou UDs and disturbance increased during spring and fall following relatively cold, snowy winters (high NPI), but overall, climate did not explain changes in caribou distribution over time. We provide evidence supporting the hypothesis that caribou populations adjust their spatial distribution in relation to anthropogenic landscape change. Our findings could have implications for population persistence if distributional shifts result in greater use of alpine habitat during winter. Monitoring long‐term changes in the distribution of populations is a valuable component of conservation planning for species at risk in disturbed landscapes.     

A spatial theory for characterizing predator–multiprey interactions in heterogeneous landscapes

Trophic interactions in multiprey systems can be largely determined by prey distributions. Yet, classic predator–prey models assume spatially homogeneous interactions between predators and prey. We developed a spatially informed theory that predicts how habitat heterogeneity alters the landscape-scale distribution of mortality risk of prey from predation, and hence the nature of predator interactions in multiprey systems. The theoretical model is a spatially explicit, multiprey functional response in which species-specific advection–diffusion models account for the response of individual prey to habitat edges. The model demonstrates that distinct responses of alternative prey species can alter the consequences of conspecific aggregation, from increasing safety to increasing predation risk. Observations of threatened boreal caribou, moose and grey wolf interacting over 378 181 km² of human-managed boreal forest support this principle. This empirically supported theory demonstrates how distinct responses of apparent competitors to landscape heterogeneity, including to human disturbances, can reverse density dependence in fitness correlates.