Functional Responses In Habitat Selection Research Articles

Carnivore space use behaviors reveal variation in responses to human land modification

BackgroundSpatial behavior, including home-ranging behaviors, habitat selection, and movement, can be extremely informative in estimating how animals respond to landscape heterogeneity. Responses in these spatial behaviors to features such as human land modification and resources can highlight a species’ spatial strategy to maximize fitness and minimize mortality. These strategies can vary on spatial, temporal, and individual scales, and the combination of behaviors on these scales can lead to very different strategies among species.MethodsHarnessing the variation present at these scales, we characterized how species may respond to stimuli in their environments ranging from broad- to fine-scale spatial responses to human modification in their environment. Using 15 bobcat-years and 31 coyote-years of GPS data from individuals inhabiting a landscape encompassing a range of human land modification, we evaluated the complexity of both species’ responses to human modification on the landscape through their home range size, habitat selection, and functional response behaviors, accounting for annual, seasonal, and diel variation.ResultsBobcats and coyotes used different strategies in response to human modification in their home ranges, with bobcats broadly expanding their home range with increases in human modification and displaying temporal consistency in functional response in habitat selection across both season and time of day. Meanwhile, coyotes did not expand their home ranges with increased human modification, but instead demonstrated fine-scale responses to human modification with habitat selection strategies that sometimes varied by time of day and season, paired with functional responses in selection behaviors.ConclusionsThese differences in response to habitat, resources, and human modification between the two species highlighted the variation in spatial behaviors animals can use to exist in anthropogenic environments. Categorizing animal spatial behavior based on these spatiotemporal responses and individual variation can help in predicting how a species will respond to future change based on their current spatial behavior.

Functional responses in habitat selection as a management tool to evaluate agri-environment schemes for farmland birds

Agri-environment schemes (AES), as part of the European Union's Common Agricultural Policy, are intended to help prevent the decline of farmland biodiversity. Nevertheless, the ecological effectiveness of AES in supporting farmland bird populations remains inconclusive across studies. This inconsistency highlights a research gap: What behavioral mechanisms contribute to the variation in farmland bird populations? This variability may arise because farmland birds alter their habitat selection in response to available habitat—a phenomenon known as functional responses in habitat selection. Here, we examined the effects of AES and non-AES land-use variables on habitat selection of farmland birds, taking into account the species-specific functional response to availability. We built two types of hierarchical distance sampling models to analyze observational data of four farmland bird species from line-transect surveys during the breeding season in Saxony, Germany. First, we built mixed-effects models to estimate the marginal effects of AES and non-AES land-use variables on the occurrence of farmland birds. Second, we integrated linear models into the distance sampling model to relate habitat selection estimates to habitat availability. Results from the first mixed-effects model showed positive and negative effects of AES on habitat selection. In the second model, we observed inverse relationships between habitat selection and availability for both AES and non-AES variables. These results support the hypothesis of negative functional responses, as we found a decrease in the tendency of farmland birds to select both AES and non-AES land uses as their availability increased. Our findings suggest that the varying effects of AES on bird occurrences reported in the literature may depend on cross-study differences in AES availability. We propose that functional responses in habitat selection should be considered as a phenomenon in future AES research. Our study also highlights the importance of optimal AES provision and their spatial allocation in the agricultural landscape.

Revealing functional responses in habitat selection of rocky features and rugged terrain by Eurasian lynx (Lynx lynx) using LiDAR data

ContextMany carnivores are attracted to rugged terrain, rocky areas, and conspicuous relief features. However, most of the previous research is limited to general topographical habitat characteristics and rarely consider the effects of microhabitat characteristics.ObjectivesWe used the Eurasian lynx (Lynx lynx) as a model species to investigate the effects of microhabitat characteristics and human infrastructure on habitat selection. We also tested whether there is evidence for a functional response in habitat selection across a large gradient of habitat availability.MethodsWe developed a new approach for detecting rocky outcrops from airborne LiDAR data. In combination with other remote sensing techniques and GPS-telemetry data, we assessed lynx habitat selection and functional responses across two geologically contrasting areas in Europe.ResultsWe detected > 1 million rocky outcrops and confirmed their strong selection by lynx. Lynx also selected steep, rugged, and rocky areas, especially for day-resting sites. Furthermore, lynx avoided paths during the day but selected them and other linear anthropogenic infrastructure during the night, indicating the behaviour-specific impact of human infrastructure. We also observed a functional response in the selection of rocky and rugged areas, as lynx’ selection of such habitats increased with their lower availability. This highlights the importance of preserving such terrains, especially when they are rare in a landscape.ConclusionsOur results highlight the importance of incorporating remote sensing techniques and data on microhabitat features in animal habitat selection research. We also recommend caution when developing new infrastructure for human recreation or promoting its use near geomorphological features and in rugged terrain.

Functional responses in American marten habitat selection indicate cumulative effects of progressive habitat change

AbstractShifts in habitat selection may serve as behavioral indicators of changing habitat quality and can provide an insight into the effects of chronic disturbance on wildlife populations before rates of occurrence and other metrics related to population viability are affected. Long‐term studies can provide unique opportunities to understand animals' behavioral responses to the cumulative effects of habitat disturbance that may be missed by short‐term studies or space‐for‐time substitution. Using the American marten (Martes americana) as a focal species, we evaluated the effects of progressive habitat disturbance on an animal population by identifying changes in habitat availability and selection over time. Marten are strongly associated with mature forest conditions and are thus sensitive to forest harvesting, which is a chronic and prevalent form of disturbance across the species' range. We developed resource selection functions that characterized both temporally consistent patterns and functional responses in the relative selection of habitats by marten at the patch scale. We used a combination of marten location data collected during three periods over 30 years of extensive forest change in northern Maine and a habitat classification scheme informed by species' behavior and designed to capture habitat changes over time. Marten increased their selection of tall (>12 m mean tree height) mature forest, including uncut forest and regenerated clearcuts, in response to decreased availability. Marten also increased avoidance of early‐successional forest (<9 m mean tree height) in response to increased availability. These functional responses in habitat selection appear to enable marten to maintain access to required resources in mature forest and limit exposure to risky or resource‐poor habitat within home ranges. However, a declining trend in the occurrence of resident adult females may indicate a limit to the functional response strategies for individuals unable to incur the energetic or prey‐related costs that may be associated with strong functional responses. Our results suggest that, despite adaptive habitat selection strategies, there are minimum thresholds of mature forest availability below which resident adults, particularly females, will struggle to persist. We recommend forest management strategies targeted at minimizing the strength of functional responses displayed by marten within occupied landscapes.

Risky business: How an herbivore navigates spatiotemporal aspects of risk from competitors and predators.

Understanding factors that influence animal behavior is central to ecology. Basic principles of animal ecology imply that individuals should seek to maximize survival and reproduction, which means carefully weighing risk against reward. Decisions become increasingly complex and constrained, however, when risk is spatiotemporally variable. We advance a growing body of work in predator-prey behavior by evaluating novel questions where a prey species is confronted with multiple predators and a potential competitor. We tested how fine-scale behavior of female mule deer (Odocoileus hemionus) during the reproductive season shifted depending upon spatial and temporal variation in risk from predators and a potential competitor. We expected female deer to avoid areas of high risk when movement activity of predators and a competitor were high. We used GPS data collected from 76 adult female mule deer, 35 adult female elk, 33 adult coyotes, and six adult mountain lions. Counter to our expectations, female deer exhibited selection for multiple risk factors, however, selection for risk was dampened by the exposure to risk within home ranges of female deer, producing a functional response in habitat selection. Furthermore, temporal variation in movement activity of predators and elk across the diel cycle did not result in a shift in movement activity by female deer. Instead, the average level of risk within their home range was the predominant factor modulating the response to risk by female deer. Our results counter prevailing hypotheses of how large herbivores navigate risky landscapes and emphasize the importance of accounting for the local environment when identifying effects of risk on animal behavior. Moreover, our findings highlight additional behavioral mechanisms used by large herbivores to mitigate multiple sources of predation and potential competitive interactions.

Individual differences in habitat selection mediate landscape level predictions of a functional response.

Predicting future space use by animals requires models that consider both habitat availability and individual differences in habitat selection. The functional response in habitat selection posits animals adjust their habitat selection to availability, but population-level responses to availability may differ from individual responses. Generalized functional response (GFR) models account for functional responses by including fixed effect interactions between habitat availability and selection. Population-level resource selection functions instead account for individual selection responses to availability with random effects. We compared predictive performance of both approaches using a functional response in elk (Cervus canadensis) selection for mixed forest in response to road proximity, and avoidance of roads in response to mixed forest availability. We also investigated how performance changed when individuals responded differently to availability from the rest of the population. Individual variation in road avoidance decreased performance of both models (random effects: β = 0.69, 95% CI 0.47, 0.91; GFR: β = 0.38, 95% CI 0.05, 0.71). Changes in individual road and forest availability affected performance of neither model, suggesting individual responses to availability different from the functional response mediated performance. We also found that overall, both models performed similarly for predicting mixed forest selection (F1, 58 = 0.14, p = 0.71) and road avoidance (F1, 58 = 0.28, p = 0.60). GFR estimates were slightly better, but its larger number of covariates produced greater variance than the random effects model. Given this bias-variance trade-off, we conclude that neither model performs better for future space use predictions.

Holling meets habitat selection: functional response of large herbivores revisited

BackgroundHolling (Can Entomol 91(5):293–320, 1959) was the first to describe a functional response between a predator’s consumption-rate and the density of its prey. The same concept can be applied to the habitat selection of herbivores, specifically, the change in relative habitat use with the change in habitat availability. Functional responses in habitat selection at a home-range scale have been reported for several large herbivores. However, a link to Holling’s original functional response types has never been drawn, although it could replace the current phenomenological view with a more mechanistically based understanding of functional responses.MethodsIn this study, discrete choice models were implemented as mixed-effects baseline-category logit models to analyze the variation in habitat selection of a large herbivore at seasonal and diurnal scales. Thus, changes in the use of land cover types with respect to their availability were investigated by monitoring 11 land cover types commonly used by roe deer (Capreolus capreolus) in the Bavarian Forest National Park, Germany. Functional response curves were then fitted using Holling’s formulas.ResultsStrong evidence of non-linear functional responses was obtained for almost all of the examined land cover types. The shape of the functional response curves varied depending on the season, the time of day, and in some cases between sexes. These responses could be referenced to Holling’s types, with a predominance of type II.ConclusionsOur results indicate that Holling’s types can be applied to describe general patterns of the habitat selection behavior of herbivores. Functional responses in habitat selection may occur in situations requiring a trade-off in the selection of land cover types offering different resources, such as due to the temporally varying physiological needs of herbivores. Moreover, two associated parameters defining the curves (prey density and predation rate) can aid in the identification of temporal variations and in determinations of the strength of the cost-benefit ratio for a specific land cover type. Application of our novel approach, using Holling’s equations to describe functional responses in the habitat selection of herbivores, will allow the assignment of general land cover attraction values, independent of availability, thus facilitating the identification of suitable habitats.

Wolf habitat selection in relation to recreational structures in a national park

AbstractAlthough most predators usually avoid human activity, some individuals instead will habituate to it. Habituation to human presence and infrastructure by predator species such as wolves may lead to conflicts implicating serious risks for public safety and for the survival of the animals involved. Accordingly, this research project aims to shed light on the relationship between wolves and recreational structures using telemetry data from 10 wolves located in the Parc National du Mont-Tremblant (Québec, Canada) and its surrounding area. Using resource selection functions (RSFs), we observed wolf habitat selection in relation to these structures during three biological periods (denning: May–June; rendezvous: June–October; and nomadic: October–April). Our results revealed that wolves selected proximity to linear structures (roads and trails) during the denning and rendezvous periods, but this selection depended on the density of such structures in the surroundings (i.e., functional response in habitat selection): wolves selected proximity to linear structures when these structures were present at greater densities. Wolves avoided housing structures (campsites, cabins, park facilities), especially when these structures were present at greater densities, suggesting that wolves perceived them as a risk. These results suggest that conflicts between visitors and wolves were unlikely to occur in campgrounds during the time of our study. This could indicate that the management measures implemented by the park following the past episodes of conflict were effective. However, wolves’ use of linear structures could lead to increased tolerance to human proximity if left unmanaged.

The density of anthropogenic features explains seasonal and behaviour-based functional responses in selection of linear features by a social predator

Anthropogenic linear features facilitate access and travel efficiency for predators, and can influence predator distribution and encounter rates with prey. We used GPS collar data from eight wolf packs and characteristics of seismic lines to investigate whether ease-of-travel or access to areas presumed to be preferred by prey best explained seasonal selection patterns of wolves near seismic lines, and whether the density of anthropogenic features led to functional responses in habitat selection. At a broad scale, wolves showed evidence of habitat-driven functional responses by exhibiting greater selection for areas near low-vegetation height seismic lines in areas with low densities of anthropogenic features. We highlight the importance of considering landscape heterogeneity and habitat characteristics, and the functional response in habitat selection when investigating seasonal behaviour-based selection patterns. Our results support behaviour in line with search for primary prey during summer and fall, and ease-of-travel during spring, while patterns of selection during winter aligned best with ease-of-travel for the less-industrialized foothills landscape, and with search for primary prey in the more-industrialized boreal landscape. These results highlight that time-sensitive restoration actions on anthropogenic features can affect the probability of overlap between predators and threatened prey within different landscapes.

Functional response of wolves to human development across boreal North America.

AimThe influence of humans on large carnivores, including wolves, is a worldwide conservation concern. In addition, human‐caused changes in carnivore density and distribution might have impacts on prey and, indirectly, on vegetation. We therefore tested wolf responses to infrastructure related to natural resource development (i.e., human footprint).LocationOur study provides one of the most extensive assessments of how predators like wolves select habitat in response to various degrees of footprint across boreal ecosystems encompassing over a million square kilometers of Canada.MethodsWe deployed GPS‐collars on 172 wolves, monitored movements and used a generalized functional response (GFR) model of resource selection. A functional response in habitat selection occurs when selection varies as a function of the availability of that habitat. GFRs can clarify how human‐induced habitat changes are influencing wildlife across large, diverse landscapes.ResultsWolves displayed a functional response to footprint. Wolves were more likely to select forest harvest cutblocks in regions with higher cutblock density (i.e., a positive functional response to high‐quality habitats for ungulate prey) and to select for higher road density in regions where road density was high (i.e., a positive functional response to human‐created travel routes). Wolves were more likely to use cutblocks in habitats with low road densities, and more likely to use roads in habitats with low cutblock densities, except in winter when wolves were more likely to use roads regardless of cutblock density.Main conclusionsThese interactions suggest that wolves trade‐off among human‐impacted habitats, and adaptively switch from using roads to facilitate movement (while also risking encounters with humans), to using cutblocks that may have higher ungulate densities. We recommend that conservation managers consider the contextual and interacting effects of footprints when assessing impacts on carnivores. These effects likely have indirect impacts on ecosystems too, including on prey species.

Revisiting the functional response in habitat selection for large herbivores: a matter of spatial variation in resource distribution?

AbstractMost habitats are distributed heterogeneously in space, forcing animals to move according to both habitat characteristics and their needs for energy and safety. Animal space use should therefore vary according to habitat characteristics, a process known as the “functional response” in habitat selection. This response has often been tested vis-à-vis the proportion of a habitat category within areas available to individuals. Measuring sought-after resources in landscape where they are continuously distributed is a challenge and we posit here that both the mean availability of a resource and its spatial variation should be measured. Accordingly, we tested for a functional response in habitat selection according to these two descriptors of the resource available for a mountain herbivore. We hypothesized that selection should decrease with mean value of resources available and increase with its spatial variation. Based on GPS data from 50 chamois females and data on the actual foodscape (i.e., distribution of edible-only biomass in the landscape), we estimated individual selection ratio (during summer months) for biomass at the home range level, comparing edible biomass in individual home ranges and the mean and standard deviation of edible biomass in their available range. Chamois being a group-living species, available accessible ranges were shared by several individuals that formed socio-spatial groups (clusters) in the population. As expected, selection ratios increased with the standard deviation of edible resources in each cluster, but unlike our prediction, was unrelated to its mean. Selection of areas richer in resources hence did not fade away when more resources were available on average, a result that may be explained by the need for this capital breeder species to accumulate fat-reserve at a high rate during summer months. Low spatial variation could limit the selection of chamois, which highlights the importance of resource distribution in the process of habitat selection.

Functional responses in habitat selection: clarifying hypotheses and interpretations

A fundamental challenge in habitat ecology and management is understanding the mechanisms generating animal distributions. Studies of habitat selection provide a lens into such mechanisms, but are often limited by unrealistic assumptions. For example, most studies assume that habitat selection is constant with respect to the availability of resources, such that habitat use remains proportional to availability. To the contrary, a growing body of work has shown the fallacy of this assumption, indicating that animals modify their behavior depending on the context at broader scales. This has been termed a functional response in habitat selection. Furthermore, a diversity of methods is employed to model functional responses in habitat selection, with little attention to how methodology might affect scientific and conservation conclusions. Here, we first review the conceptual and statistical foundations of methods currently used to model functional responses and clarify the ecological tests evaluated within each approach. We then use a combination of simulated and empirical data sets to evaluate the similarities and differences among approaches. Importantly, we identified multiple statistical issues with the most widely applied approaches to understand functional responses, including: (1) a complex and important role of random- or individual-level intercepts in adjusting individual-level regression coefficients as resource availability changes and (2) a sensitivity of results to poorly informed individual-level coefficients estimated for animals with low availability of a given resource. Consequently, we provide guidance on applying approaches that are insensitive to these issues with the goal of advancing our understanding of animal habitat ecology and management. Finally, we characterize the management implications of assuming similarity between the current approaches to model functional responses with two empirical data sets of federally threatened species: Canada lynx (Lynx canadensis) in the United States and woodland caribou (Rangifer tarandus caribou) in Canada. Collectively, our assessment helps clarify the similarities and differences among current approaches and, therefore, assists the integration of functional responses into the mainstream of habitat ecology and management.

Functional responses in animal movement explain spatial heterogeneity in animal-habitat relationships.

Understanding why heterogeneity exists in animal-habitat spatial relationships is critical for identifying the drivers of animal distributions. Functional responses in habitat selection - whereby animals adjust their habitat selection depending on habitat availability - are useful for describing animal-habitat spatial heterogeneity. However, they could be yielded by different movement tactics, involving contrasting interspecific interactions. Identifying functional responses in animal movement, rather than in emergent spatial patterns like habitat selection, could disentangle the effects of different movement behaviours on spatial heterogeneity in animal-habitat relationships. This would clarify how functional responses in habitat selection emerge and provide a general tool for understanding the mechanistic drivers of animal distributions. We tested this approach using data from GPS-collared woodland caribou (Rangifer tarandus), a prey species under top-down control. We tested how caribou selected and moved with respect to a key resource (lichen-conifer stands) as a function of the availability of surrounding refuge land-cover (closed-conifer stands), using step selection functions. Caribou selected resource patches more strongly in areas richer in refuge land-cover - a functional response in habitat selection. However, adjustments in multiple movement behaviours could have generated this pattern: stronger directed movement towards resource patches and/or longer residency within resource patches, in areas richer in refuges. Different contributions of these behaviours would produce contrasting forager spatial dynamics. We identified functional responses in both movement behaviours: caribou were more likely to move towards resource patches in areas richer in refuge land-cover, and to remain in these patches during movement steps. This tactic enables caribou to forage for longer in safer areas where they can rapidly seek refuge in dense cover when predators are detected. Our study shows that functional responses in movement can expose the context-dependent movement decisions that generate heterogeneity in animal-habitat spatial relationships. We used these functional responses to characterise anti-predator movement tactics employed by a large herbivore, but they could be applied in many different scenarios. The movement rules from functional responses in movement are well-suited to integration in spatial explicit individual-based models for forecasting animal distributions in landscapes undergoing environmental change.

Habitat functional response mitigates reduced foraging opportunity: implications for animal fitness and space use

Animals selectively use landscapes to meet their energetic needs, and trade-offs in habitat use may depend on availability and environmental conditions. For example, habitat selection at high temperatures may favor thermal cover at the cost of reduced foraging efficiency under consistently warm conditions. Our objective was to examine habitat selection and space use in distinct populations of moose (Alces alces). Hypothesizing that endotherm fitness is constrained by heat dissipation efficiency, we predicted that southerly populations would exhibit greater selection for thermal cover and reduced selection for foraging habitat. We estimated individual step selection functions with shrinkage for 134 adult female moose in Minnesota, USA, and 64 in Ontario, Canada, to assess habitat selection with variation in temperature, time of day, and habitat availability. We averaged model coefficients within each site to quantify selection strength for habitats differing in forage availability and thermal cover. Moose in Ontario favored deciduous and mixedwood forest, indicating selection for foraging habitat across both diel and temperature. Habitat selection patterns of moose in Minnesota were more dynamic and indicated time- and temperature-dependent trade-offs between use of foraging habitat and thermal cover. We detected a scale-dependent functional response in habitat selection driven by the trade-off between selection for foraging habitat and thermal cover. Landscape composition and internal state interact to produce complex patterns of space use, and animals exposed to increasingly high temperatures may mitigate fitness losses from reduced foraging efficiency by increasing selection for foraging habitat in sub-prime foraging landscapes.

Quantifying consistent individual differences in habitat selection.

Habitat selection is a fundamental behaviour that links individuals to the resources required for survival and reproduction. Although natural selection acts on an individual's phenotype, research on habitat selection often pools inter-individual patterns to provide inferences on the population scale. Here, we expanded a traditional approach of quantifying habitat selection at the individual level to explore the potential for consistent individual differences of habitat selection. We used random coefficients in resource selection functions (RSFs) and repeatability estimates to test for variability in habitat selection. We applied our method to a detailed dataset of GPS relocations of brown bears (Ursus arctos) taken over a period of 6years, and assessed whether they displayed repeatable individual differences in habitat selection toward two habitat types: bogs and recent timber-harvest cut blocks. In our analyses, we controlled for the availability of habitat, i.e. the functional response in habitat selection. Repeatability estimates of habitat selection toward bogs and cut blocks were 0.304 and 0.420, respectively. Therefore, 30.4 and 42.0% of the population-scale habitat selection variability for bogs and cut blocks, respectively, was due to differences among individuals, suggesting that consistent individual variation in habitat selection exists in brown bears. Using simulations, we posit that repeatability values of habitat selection are not related to the value and significance of β estimates in RSFs. Although individual differences in habitat selection could be the results of non-exclusive factors, our results illustrate the evolutionary potential of habitat selection.

Grain-dependent functional responses in habitat selection

Context Spatial scale is an important consideration for understanding how animals select habitat, and multi-scalar designs in resource selection studies have become increasingly common. Despite this, examination of functional responses in habitat selection at multiple scales is rare. The perceptual range of an animal changes as a function of vegetation association, suggesting that use, selection and functional responses may all be habitat- and scale-dependent.

Functional responses in habitat selection are density dependent in a large herbivore

Habitat selection is complex due to density dependence and functional responses, defined as variation in relative habitat use depending on availability. In this study we unite these concepts by empirically testing for density‐dependent functional responses in habitat selection using a large herbivore, elk Cervus canadensis manitobensis, as a model species. Theory on density‐dependent habitat selection predicts specialised behaviour when densities are low with a gradual switch to generalist behaviour (more even selection of habitat) as competition intensifies. This suggests that functional responses in habitat selection should be positive when competition is low, but that density may have a negative effect on the functional response due to constraining effects of competition on habitat use and availability. We tested this prediction using data from Global Positioning System (GPS) collared elk (n = 33) and empirical data on spatiotemporal variation in local density during winter in Riding Mountain National Park, Manitoba, Canada (2002–2011). As local density increased, winter home range size decreased and the proportion of mixed forest (providing shelter and forage to elk) used and available within the home range also decreased. Our resource selection function (RSF) revealed clear density‐dependent effects in selection, being strongest (or weakest) for high quality (or low quality) habitat types at lowest observed density leading to more even selection as densities increased. The functional response in mixed forest selection was negatively affected by local density. Increasing availability of mixed forest in the home range was associated with higher selection at low density (positive functional response); no effect of availability on selection at moderate density (no functional response); and lower selection as availability increased at high density (negative functional response). Our study demonstrates that this process is largely driven by the negative effect of density on home range size as it constrains use and availability of habitat.

Adjustments in habitat selection to changing availability induce fitness costs for a threatened ungulate

Summary Functional responses in habitat selection occur when individuals adjust their selection of habitat features as a function of the availability of those features. Functional responses in habitat selection are generally assumed to be fitness‐rewarding tactics and are used to guide conservation actions. Fitness consequences of functional responses, however, have rarely been evaluated. Eighty‐three caribou were followed with GPS collars to establish the link between functional responses in habitat selection and adult female survival, a strong fitness correlate for caribou. We measured how caribou avoidance of mixed/deciduous stands and 6‐ to 20‐year‐old clear‐cuts varied with the proportion of 6‐ to 20‐year‐old clear‐cuts within their 100% minimal convex polygon (MCP), and if these functional responses were linked to survival. Mixed/deciduous stands and 6‐ to 20‐year‐old clear‐cuts are risky for caribou because they are selected by moose, thereby attracting wolves and increasing predation risk for caribou. Caribou avoided mixed/deciduous stands, especially when 6‐ to 20‐year‐old clear‐cuts comprised a large proportion of their MCP, but this functional response did not differ between caribou that died and those that survived. When the proportion of 6‐to 20‐year‐old clear‐cuts in the MCP was low, caribou generally had low odds of occurring near 6‐ to 20‐year‐old clear‐cuts. However, when the proportion of clear‐cuts in the MCP was relatively high, caribou that strongly increased their odds of being near 6‐ to 20‐year‐old clear‐cuts were generally those that died. Synthesis and applications. Assessing the fitness consequences of how animals respond to habitat disturbances is central to wildlife conservation. We demonstrate that distinct functional responses in habitat selection involve different mortality risks and that population dynamics should depend on the frequency of the different tactics observed within populations. Individuals that persistently select riskier areas should be important drivers of population decline. Caribou mortality could be reduced by decreasing the appeal of 6‐ to 20‐year‐old clear‐cuts for moose by removing deciduous vegetation through cleaning, which should reduce the selection of wolves for these stands. Removing deciduous vegetation should be especially effective in areas where those clear‐cuts comprise a large proportion of the landscape, because this is where a subset of the caribou population experiences high mortality rates by selecting 6‐ to 20‐year‐old clear‐cuts.

Density‐dependent functional responses in habitat selection by two hosts of the raccoon rabies virus variant

Spatio‐temporal variations in conspecific density and resource availability are two of the main factors responsible for plasticity in habitat selection. Despite the need for habitat selection models that can accurately predict animal distribution given the plasticity in the selection process, no study has assessed the synergistic effects of these factors on habitat selection. We investigated density‐dependent functional responses by raccoons (Procyon lotor) and striped skunks (Mephitis mephitis), two of the main hosts of the rabies virus in North America. We monitored 54 raccoons and 12 striped skunks with Global‐Positioning‐System collars in a landscape dominated by corn fields and forest patches. We built resource selection functions to evaluate if the selection of corn fields varied with conspecific density and corn field availability within 100% minimum convex polygons. Raccoons altered their selection of corn fields depending on both conspecific density and corn‐forest edge density or corn field proportion. In areas of low corn‐forest edge densities and a low corn field proportion, raccoons showed stronger selection for corn fields when few conspecifics were present. At high conspecific densities, the selection of corn fields was stronger in areas with high corn‐forest edge densities and a low corn field proportion. For striped skunks, we did not detect any synergistic effect of density‐dependence and functional responses. Unlike raccoons, striped skunks displayed a selection that was strongest for agricultural corridors. We show that functional responses in habitat selection can be density‐dependent. In a context of infectious disease dynamics, modeling density‐dependence in functional responses increases the ability to predict spatio‐temporal variations in the distribution of reservoir species and thus, to delineate areas at high animal densities where the risk of disease outbreaks is relatively high. For example, the omission of density‐dependence in functional responses underestimated the relative probability of raccoon occurrence in corn fields, while overestimating the relative probability of occurrence in anthropogenic areas and wetlands. Our study underscores the relevance of considering the complexity of habitat selection by all hosts of a zoonosis. Cost‐effective control and prevention programs used to limit disease spread can benefit from accounting for density‐dependent functional responses of a multi‐host disease system.

Flexible habitat selection by cougars in response to anthropogenic development

Large carnivore populations are imperiled worldwide through habitat loss, insufficient prey, and intolerance by people, promoting concerns that their populations will persist only in protected areas and remote regions. This assumption is challenged by recent colonization of areas with high human use in the former ranges of several carnivore species, including cougars (Puma concolor). We hypothesized that cougars adjust their behavior to accommodate anthropogenic development, facilitating successful use of modified habitats. We tested this hypothesis using resource selection functions for 42 individual cougars maintaining home ranges across a gradient of anthropogenic development. We evaluated variation in selection among individual cougars by time of day, home-range location (i.e., rural vs. wilderness cougars), and using functional response curves. Cougars stayed further away from both roads and buildings during the day than they did at night, and wilderness cougars showed stronger avoidance of anthropogenic features than did their rural counterparts. Selection patterns of individual cougars varied greatly, and some of this variation was explained by the functional response. For example, cougars decreased their avoidance of some anthropogenic features as those features became more prevalent on the landscape. Failure to account for potential functional responses in habitat selection could lead to overestimation of negative impacts of development for adaptable large carnivores. Given high adaptability to anthropogenic habitat modification displayed by cougars, cougar persistence in increasingly developed landscapes may depend more on human tolerance than on maintaining large tracts of pristine wilderness.