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.

Outdoor recreation is increasingly recognized to impact nature and wildlife, yet few studies have examined recreation within large natural landscapes that are critical habitat to some of our most rare and potentially disturbance‐sensitive species. Over six winters (2010–2015) and four study areas (>1.1 million ha) in Idaho, Wyoming, and Montana, we studied the responses of wolverines (Gulo gulo) to backcountry winter recreation. We fit Global Positioning System (GPS) collars to 24 individual wolverines and acquired >54,000 GPS locations over 39 animal‐years during winter (January–April). Simultaneously, we monitored winter recreation, collecting ~6000 GPS tracks (~200,000 km) from backcountry recreationists. We combined the GPS tracks with trail use counts and aerial recreation surveys to map the extent and relative intensity of motorized and non‐motorized recreation. We integrated our wolverine and backcountry recreation data to (1) assess patterns of wolverine habitat selection and (2) evaluate the effect of backcountry recreation on wolverine habitat relationships. We used resource selection functions to model habitat selection of male and female wolverines within their home ranges. We first modeled habitat selection for environmental covariates to understand male and female habitat use then incorporated winter recreation covariates. We assessed the potential for indirect habitat loss from winter recreation and tested for functional responses of wolverines to differing levels and types of recreation. Motorized recreation occurred at higher intensity across a larger footprint than non‐motorized recreation in most wolverine home ranges. Wolverines avoided areas of both motorized and non‐motorized winter recreation with off‐road recreation eliciting a stronger response than road‐based recreation. Female wolverines exhibited stronger avoidance of off‐road motorized recreation and experienced higher indirect habitat loss than male wolverines. Wolverines showed negative functional responses to the level of recreation exposure within the home range, with female wolverines showing the strongest functional response to motorized winter recreation. We suggest indirect habitat loss, particularly to females, could be of concern in areas with higher recreation levels. We speculate that the potential for backcountry winter recreation to affect wolverines may increase under climate change if reduced snow pack concentrates winter recreationists and wolverines in the remaining areas of persistent snow cover.

Ecologists and managers are motivated to predict the distribution of animals across landscapes as well as understand the mechanisms giving rise to that distribution. Satisfying this motivation requires an integrated framework that characterizes multi‐scale habitat use and selection, as well as builds predictive models such as resource selection functions. However, the assumption of constant habitat use or selection is often made in such analyses, which ignores the possibility that individuals experiencing different conditions might respond differently. Assessing functional responses in habitat use evaluates how animal behavior changes with differing environmental conditions, which has basic and applied utility. Here, we combined these ideas into an integrated process that characterizes habitat relationships, predicts habitat, and assesses behavioral differences with changing environmental conditions. Our species of interest was Canada lynx (Lynx canadensis) in the Northern Rocky Mountains, which is a rare and federally threatened forest carnivore. Through our process, we developed multi‐scale predictions of lynx distribution and learned that across scales and seasons, lynx use more mature, spruce‐fir forests than any other structure stage or species. Intermediate snow depths and the distribution of snowshoe hares (Lepus americanus) were the strongest predictors of where lynx selected their home ranges. Within their home ranges, female and male lynx increasingly used advanced regeneration forest structures as they became more available (up to a maximum availability of 40%). These patterns supported the bottom‐up mechanisms regulating Canada lynx in that advanced regeneration generally provides the most abundant snowshoe hares, while mature forest is where lynx appear to hunt efficiently. However, lynx exhibited decreasing use of stand initiation structures (up to a maximum availability of 25%). Land managers have an opportunity to promote lynx habitat in the form of advanced regeneration, but are required to go through the stand initiation phase. Thus, managers can apply the relative proportions of forest structure classes along with our response curves to inform landscape actions (e.g., timber harvest) targeted at facilitating the forest mosaic used and selected by Canada lynx. Collectively, the insights gleaned from our approach advance habitat conservation efforts and consequently are of broad utility to applied ecologists and managers.

Sagebrush (Artemisia tridentata) ecosystems are declining due to biological invasions and changes in fire regimes. Understanding how ecosystem changes influence functionally important animals such as ecosystem engineers is essential to conserve ecological functions. American badgers (Taxidea taxus) are an apex predator and ecosystem engineer in that they redistribute large amounts of soil within sagebrush ecosystems. Piute ground squirrels (Urocitellus mollis) are also an ecosystem engineer as well as an essential prey resource for many predators, including badgers. Our research objective was to evaluate the relative importance of biological invasions and fire, abiotic factors, and biotic factors on badgers and ground squirrels. We sampled 163 1‐ha plots during April‐June across a gradient of burn histories within a 1962 km² study area in southern Idaho, USA. At each plot, we characterized occupancy of ground squirrels and badgers and relative abundance of ground squirrels. Additionally, we characterized soil texture, climate, connectivity and dispersal potential, fire frequency, grazing, and cover of many plant species including a highly invasive exotic annual grass (cheatgrass; Bromus tectorum). We used an integrated approach to evaluate competing hypotheses concerning factors influencing occupancy and abundance. Results suggested that occupancy of ground squirrels was positively associated with long‐term precipitation, dispersal potential, and fine‐grained soil. Abundance of ground squirrels was positively associated with fine‐grained soil, but negatively associated with cheatgrass, fire frequency, agriculture, and shrub cover. Badger occupancy was positively associated with ground squirrel occupancy and agriculture, which indicated affinity to prey. Our results provide insight into the relative influence of abiotic and biotic factors on predator and prey, and highlight how effects change across different population parameters. Our research suggests that widespread environmental change within sagebrush ecosystems, especially the fire‐cheatgrass cycle (e.g., invasion of cheatgrass and increased fire frequency) and human land disturbances, are directly and indirectly influencing ground squirrels and badgers. However, we also found evidence that efforts to mitigate these stressors, for example, establishing bunchgrasses postfire, may provide targeted conservation strategies that promote these species and thus preserve the burrowing and trophic functions they provide.

Understanding intrinsic and extrinsic drivers of reproductive success is central to advancing animal ecology and characterizing critical habitat. Unfortunately, much of the work examining drivers of reproductive success is biased toward particular groups of organisms (e.g., colonial birds, large herbivores, capital breeders). Long‐lived mammalian carnivores that are of conservation concern, solitary, and territorial present an excellent situation to examine intrinsic and extrinsic drivers of reproductive success, yet they have received little attention. Here, we used a Canada lynx (Lynx canadensis) data set, from the southern periphery of their range, to determine if reproductive success in a solitary carnivore was consistent with capital or income breeding. We radio‐marked and monitored 36 female Canada lynx for 98 lynx years. We evaluated how maternal characteristics and indices of food supply (via forest structure) in core areas influenced variation in body condition and reproductive success. We characterized body condition as mass/length and reproductive success as whether a female produced a litter of kittens for a given breeding season. Consistent with life‐history theory, we documented a positive effect of maternal age on body condition and reproductive success. In contrast to predictions of capital breeding, we observed no effect of pre‐pregnancy body condition on reproductive success in Canada lynx. However, we demonstrated statistical effects of forest structure on reproductive success in Canada lynx, consistent with predictions of income breeding. The forest characteristics that defined high success included (1) abundant and connected mature forest and (2) intermediate amounts of small‐diameter regenerating forest. These attributes are consistent with providing abundant, temporally stable, and accessible prey resources (i.e., snowshoe hares; Lepus americanus) for lynx and reinforce the bottom‐up mechanisms influencing Canada lynx populations. Collectively, our results suggest that lynx on the southern range periphery exhibit an income breeding strategy and that forest structure supplies the income important for successful reproduction. More broadly, our insights advance the understanding of carnivore ecology and serve as an important example on integrating long‐term field studies with ecological theory to improve landscape management.

Snowshoe hares (Lepus americanus) are an ecologically important herbivore because they modify vegetation through browsing and serve as a prey resource for multiple predators. We implemented a multiscale approach to characterize habitat relationships for snowshoe hares across the mixed conifer landscape of the northern Rocky Mountains,USA. Our objectives were to (1) assess the relationship between horizontal cover and snowshoe hares, (2) estimate how forest metrics vary across the gradient of snowshoe hare use and horizontal cover, and (3) model and map snowshoe hare occupancy and intensity of use. Results indicated that both occupancy and intensity of use by snowshoe hares increased with horizontal cover and that the effect became stronger as intensity of use increased. This underscores the importance of dense horizontal cover to achieve high use, and likely density, of snowshoe hares. Forest structure in areas with high snowshoe hare use and horizontal cover was characterized as multistoried with dense canopy cover and medium‐sized trees (e.g., 12.7–24.4 cm). The abundance of lodgepole pine (Pinus contorta) was associated with snowshoe hare use within a mixed conifer context, and the only species to increase in abundance with horizontal cover wasEngelmann spruce (Picea engelmannii) and subalpine fir (Abies lasiocarpa). Our landscape‐level modeling produced similar patterns in that we observed a positive effect of lodgepole pine and horizontal cover on both occupancy and use by snowshoe hares, but we also observed a positive yet parabolic effect of snow depth on snowshoe hare occupancy. This work is among the first to characterize the multiscale habitat relationships of snowshoe hares across a mixed conifer landscape as well as to map their occupancy and intensity of use. Moreover, our results provide stand‐ and landscape‐level insights that directly relate to management agencies, which aids in conservation efforts of snowshoe hares and their associated predators.