The development of forage production and utilization gradients around livestock watering points

Large herbivores can impose spatial patterns on otherwise homogeneous vegetation, but how these patterns change through time is poorly understood. Domestic livestock pastures are model systems for studying how foraging behavior influences the development of coupled grazing and vegetation patterns. We sampled forage production and utilization by cattle along distance-from-water gradients to provide a snapshot of grazing and vegetation patterns, and then evaluated the ability of simulation models to qualitatively reproduce these patterns. In the field, forage production increased with distance from water, as expected, but utilization peaked at intermediate distances from water in two of three study areas. Likewise, simulations based on a variety of foraging strategies produced gradients in forage production and, after forage availability near water declined sufficiently, peaks in utilization at intermediate distances. Distance-from-water gradients thus represent cumulative but not necessarily present day gradients in grazing intensity. The model with a foraging strategy based on time minimization produced slightly more realistic patterns in forage abundance than a model based on energy maximization, although results were sensitive to the value of the threshold for rejecting sites of low forage biomass. However, all models produced implausible thresholds in grazing and forage distribution, suggesting that factors besides resource distribution influence herbivore distributions. Moreover, different foraging rules produced similar vegetation gradients, especially on point water source landscapes, illustrating the difficulty of inferring foraging processes from vegetation patterns.     

Interactions between landscape structure and animal behavior: the roles of heterogeneously distributed resources and food deprivation on movement patterns

To examine how resource distributions affect the movement behaviors of fed and food-deprived Eleodes extricata Say darkling beetles (Coleoptera: Tenebrionidae), we experimentally manipulated the dispersion of food to create clumped, random, and uniform distributions in an otherwise homogeneous 25-m² experimental field landscape. Quantitative measures of the tortuosity, net linear displacement, overall path length, and velocity of beetle movement pathways showed that food-deprived beetles generally moved more slowly and over shorter distances than did fed beetles. This effect was mediated by the spatial distribution of food, however; food distributed randomly over the landscape evoked more tortuous paths over larger overall distances. The foraging movements of food-deprived beetles were most different from those of fed individuals in treatments with randomly distributed food resources. These results show that the influence of spatial structure on individuals depends not only on the arrangement of pattern but also on the function that the structure plays. Thus, 'spatial structure' is defined not only by physical characteristics of the landscape but also by how that structure is used by animals.     

Foraging behavior of browsing ruminants in a heterogeneous landscape

Movement patterns of white-tailed deer (Odocoileus virginianus) and Spanish goats (Carpa hircus) were mapped and analyzed to test the hypothesis that foraging movements and behaviors within an Acacia shrub community are significantly related to environmental heterogeneity. Animal response to plant community heterogeneity was characterized using foraging velocity and the animals' foraging path fractal dimension (D_d). Environmental heterogeneity was characterized using the perimeter:area fractal dimension, which represents the shape of shrubs, and the grid count fractal dimension, which represents shrub spatial distribution. The foraging paths of deer were straighter and more directed (D_d = 1.27) than those of goats (D_d = 1.53), and deer responded to the shape of shrub patches, more so than to shrub distribution. The tortuosity of goat foraging paths was apparently affected by distribution of blackbrush (Acacia rigidula) and shrubby bluesage (Salvia ballotiflora). Foraging velocity of deer was affected by the distribution and shape complexity of guajillo (A. berlandieri), which was a major dietary component. In contrast, foraging velocity of goats was affected by the shape complexity of the entire shrub community and by the distribution of ceniza (Leucophylum frutescens), a non-dietary, but prevalent component of the plant community. Results indicate that these two browsing herbivores perceive the same landscape differently.     

The effect of plant spatial pattern within a patch on foraging selectivity of grazing sheep

Plant spatial pattern has been considered as one of the most important factors influencing forage selection of herbivores in natural grasslands. Previous work has emphasized the effects of spatial distribution patterns of food resource at the scale of whole plant communities. Our objective was to explore whether changes in spatial patterns of food within a patchy site affected forage selection of sheep within and among patches. We conducted a manipulative experiment using three native plant species of different palatability and abundance to artificially create three different quality patches in each treatment. We compared the effects of aggregated and randomly dispersed patterns, within high, medium, and low quality patches respectively, on sheep forage selection. Effects of plant spatial patterns within a patch on sheep forage selection of the patch itself strongly depended on the patch quality. For high quality patches, random dispersion of food resources significantly decreased sheep consumption of the palatable plant within the patch. This effect was reversed in low quality patches, and was not significant in medium quality patches. Changes in plant spatial patterns within high quality patches greatly influenced sheep forage selection of other patches. However, changes in plant spatial patterns within medium or low quality patches significantly influenced foraging responses of sheep only for high quality patches. We therefore conclude that high quality resource sites are the most influential and susceptible foraging areas. Our results highlight the importance of high quality resource sites when considering grazing grassland conservation and management.     

Estimating landscape resistance from habitat suitability: effects of data source and nonlinearities

Context

Conservation corridors must facilitate long-distance dispersal movements to promote gene flow, prevent inbreeding, and allow animals to shift ranges with climate change. Least-cost models are used to identify areas that support long-distance movement. These models rely on estimates of landscape resistance, which are typically derived from habitat suitability.

Objectives

We examine two key steps in estimating resistance from habitat suitability: choosing a procedure to estimate habitat suitability, and choosing a transformation function to translate habitat suitability into resistance.

Methods

We used linear and nonlinear functions to convert three types of habitat suitability estimates (from expert opinion, resource selection functions, and step selection functions) into resistances for elk (Cervus canadensis) and desert bighorn sheep (Ovis canadensis nelsoni). We evaluated the resulting resistance maps on an independent set of observed long-distance, prospecting movements.

Results

A negative exponential function best described the relationship between resistance values and habitat suitability for desert bighorn sheep indicating long-distance movers readily travel through moderately-suitable areas and avoid only the least suitable habitat. For desert bighorn sheep, all three suitability estimates performed better than chance, and resource and step selection functions outperformed expert opinion. For elk, all three suitability estimates performed the same as chance.

Conclusions

When designing corridors to facilitate long-distance movements of mobile animals, we recommend transforming habitat suitability into resistance with a negative exponential function. Use of an exponential transformation means that larger fractions of the landscape offer low resistance, allowing greater flexibility in where a corridor is located.

     

Declines in moose population density at Isle Royle National Park,MI, USA and accompanied changes in landscape patterns

Ungulate herbivores create patterns of forage availability, plant species composition, and soil fertility as they range across large landscapes and consume large quantities of plant material. Over time, herbivore populations fluctuate, producing great potential for spatio-temporal landscape dynamics. In this study, we extend the spatial and temporal extent of a long-term investigation of the relationship of landscape patterns to moose foraging behavior at Isle Royale National Park, MI. We examined how patterns of browse availability and consumption, plant basal area, and soil fertility changed during a recent decline in the moose population. We used geostatistics to examine changes in the nature of spatial patterns in two valleys over 18 years and across short-range and long-range distance scales. Landscape patterns of available and consumed browse changed from either repeated patches or randomly distributed patches in 1988–1992 to random point distributions by 2007 after a recent record high peak followed by a rapid decline in the moose population. Patterns of available and consumed browse became decoupled during the moose population low, which is in contrast to coupled patterns during the earlier high moose population. Distributions of plant basal area and soil nitrogen availability also switched from repeated patches to randomly distributed patches in one valley and to random point distributions in the other valley. Rapid declines in moose population density may release vegetation and soil fertility from browsing pressure and in turn create random landscape patterns.     

Introducing spatial grazing impacts into the prediction of moorland vegetation dynamics

Grazing by large herbivores is a major determinant of vegetation dynamics in many semi-natural ecosystems, including the replacement of heather moorland by rough grassland in the British uplands. Herbivore foraging is influenced by vegetation patterns and, in turn, their grazing drives vegetation dynamics. Although vegetation impacts are local, spatially heterogeneous local impacts can have different consequences as would the same impacts distributed uniformly. We constructed a simulation model of the spatial effects of grazing by sheep on the vegetation dynamics of heather moorland, a vegetation community of international conservation importance in the UK. The model comprised three sub-models to predict (1) annual average heather utilisation, (2) spatial variation in heather utilisation (higher near the edge of grass patches) and (3) competition between heather and grass. Here we compare the predicted heather utilisation and vegetation dynamics of the spatial model, relative to those of a non-spatial model. The spatial model resulted in a reduced loss of heather cover for a given sheep stocking rate. The model demonstrates how spatial interactions between large herbivores and their forage drive vegetation dynamics, leading to changes in community structure and composition. Indeed, omitting spatial effects in grazing models may lead to inaccurate predictions. We have shown that ecosystem modelling, based around an iterative dialogue between developers and experienced researchers, has the potential to make a substantial contribution towards the conservation and management of vulnerable landscapes. Combining modelling with experimental studies will facilitate progress towards understanding long-term vegetation/herbivore dynamics.     

Introducing spatial grazing impacts into the prediction of moorland vegetation dynamics

Grazing by large herbivores is a major determinant of vegetation dynamics in many semi-natural ecosystems, including the replacement of heather moorland by rough grassland in the British uplands. Herbivore foraging is influenced by vegetation patterns and, in turn, their grazing drives vegetation dynamics. Although vegetation impacts are local, spatially heterogeneous local impacts can have different consequences as would the same impacts distributed uniformly. We constructed a simulation model of the spatial effects of grazing by sheep on the vegetation dynamics of heather moorland, a vegetation community of international conservation importance in the UK. The model comprised three sub-models to predict (1) annual average heather utilisation, (2) spatial variation in heather utilisation (higher near the edge of grass patches) and (3) competition between heather and grass. Here we compare the predicted heather utilisation and vegetation dynamics of the spatial model, relative to those of a non-spatial model. The spatial model resulted in a reduced loss of heather cover for a given sheep stocking rate. The model demonstrates how spatial interactions between large herbivores and their forage drive vegetation dynamics, leading to changes in community structure and composition. Indeed, omitting spatial effects in grazing models may lead to inaccurate predictions. We have shown that ecosystem modelling, based around an iterative dialogue between developers and experienced researchers, has the potential to make a substantial contribution towards the conservation and management of vulnerable landscapes. Combining modelling with experimental studies will facilitate progress towards understanding long-term vegetation/herbivore dynamics.     

Introducing spatial grazing impacts into the prediction of moorland vegetation dynamics

Grazing by large herbivores is a major determinant of vegetation dynamics in many semi-natural ecosystems, including ghe replacement of heather moorland by rough grassland in the British uplands. Herbivore foraging is influenced by vegetation patterns and, in turn, their grazing drives vegetation dynamics. Although vegetation impacts are local, spatially heterogeneous local impacts can have different conseqences as would the same impact distributed uniformly. We constructed a simulation model of the spatial effects of grazing by sheep on the vegetation dynamics of heather moorland, a vegetation community of international conservation importance in the UK. The model comprised three sub-models to predict (1) annual average heather utilisation, (2) spatial variation in heather utilisation (higher near the edge of grass patches) and (3) competition between heather and grass. Here we compare the predicted heather utilisation and vegetation dynamics of the spatial model, relative to those of a non-spatial model. The spatial model resulted in a reduced loss of heather cover for a given sheep stocking rate. The model demonstrtaes how spatial interactions between large herbivores and their forage drive vegetation dynamics, leading to changes in community structure and composition. Indeed, omitting spatial effects in grazing models may lead to inaccurate predictions. We have shown that ecosystem modelling, based around an iterative dialogue between developers and experienced researchers, has the potential to make a substantial contribution towards the conservation and management of vulnerable landscapes. Combining modelling with experimental studies will facilitate progress towards understanding long-term vegetation/herbivore dynamics.     

Cost analysis of remotely sensed foraging paths in patchy landscapes with plant anti-herbivore defenses (Patagonia, Argentina)

We developed metrics at a landscape scale to evaluate the costs and rewards experienced by large herbivores while foraging in natural vegetation with patchy anti-herbivore plant structures. We show an application of these metrics to the analysis of 16,000 records of positions at successive 1 min intervals of free-ranging ewes (Ovis aries) harnessed with Global-Positioning-System (GPS) loggers, in a large paddock of the Patagonian Monte shrublands (Argentina). Dominant shrubs in the area display numerous anti-herbivore defenses (spiny-resinous leaves, thorny stems, etc.) protecting them from grazing and herbivore trampling. Preferred grasses and forbs constitute a minor part of aboveground plant biomass and grow in relatively open areas among or around shrub patches. We mapped the movement speed of ewes onto high-resolution aerial photographs of the grazed paddocks and estimated costs and rewards along their paths based on algorithms of surface cost theory. Ewes explored areas of sparse vegetation at low speeds compatible with predominant grazing, and increased their speed when crossing denser shrubby patches. The cost algorithm was applied to evaluate daily searching costs as well as grazing rewards in relation to the length of daily searching paths. The observed path lengths and search speeds were consistent with those that compensate costs and rewards of the grazing activities as estimated by the surface cost analysis. We conclude that the technique presented here constitutes a valuable tool to quantify the effect of landscape characteristics on behavioral traits of grazing animals in similar environments.     

Evidence of Hierarchical Patch Dynamics in an East African savanna?

The Hierarchical Patch Dynamics Paradigm provides a conceptual framework for linking pattern, process and scale in ecosystems, but there have been few attempts to test this theory because most ecological studies focus on only one spatial scale, or are limited in their temporal scope. Here I use palaeoecological techniques (analysis of fossil pollen and stable carbon isotopes) to compare vegetation heterogeneity in an east African savanna at three spatial scales, over hundreds of years. The data show that patterns of vegetation change are different at the three spatial scales of observation, and suggest that different ecological processes dominate tree abundance at micro, local and landscape scales. Interactions between plants, disturbance (e.g., by fire and herbivores), climate and soil type may influence tree density at differing spatial and temporal scales. This hierarchical explanation of savanna vegetation dynamics could inform future biodiversity conservation and management in savannas. This revised version was published online in July 2006 with corrections to the Cover Date.     

Dispersal success on fractal landscapes: a consequence of lacunarity thresholds

Habitat fragmentation is expected to disrupt dispersal, and thus we explored how patch metrics of landscape structure, such as percolation thresholds used to define landscape connectivity, corresponded with dispersal success on neutral landscapes. We simulated dispersal as either a purely random process (random direction and random step lengths) or as an area-limited random walk (random direction, but movement limited to an adjacent cell at each dispersal step) and quantified dispersal success for 1000 individuals on random and fractal landscape maps across a range of habitat abundance and fragmentation. Dispersal success increased with the number of cells a disperser could search (m), but poor dispersers (m<5) searching via area-limited dispersal on fractal landscapes were more successful at locating suitable habitat than random dispersers on either random or fractal landscapes. Dispersal success was enhanced on fractal landscapes relative to random ones because of the greater spatial contagion of habitat. Dispersal success decreased proportionate to habitat loss for poor dispersers (m=1) on random landscapes, but exhibited an abrupt threshold at low levels of habitat abundance (p<0.1) for area-limited dispersers (m<10) on fractal landscapes. Conventional metrics of patch structure, including percolation, did not exhibit threshold behavior in the region of the dispersal threshold. A lacunarity analysis of the gap structure of landscape patterns, however, revealed a strong threshold in the variability of gap sizes at low levels of habitat abundance (p<0.1) in fractal landscapes, the same region in which abrupt declines in dispersal success were observed. The interpatch distances or gaps across which dispersers must move in search of suitable habitat should influence dispersal success, and our results suggest that there is a critical gap-size structure to fractal landscapes that interferes with the ability of dispersers to locate suitable habitat when habitat is rare. We suggest that the gap structure of landscapes is a more important determinant of dispersal than patch structure, although both are ultimately required to predict the ecological consequences of habitat fragmentation.     

Evidence of Hierarchical Patch Dynamics in an East African savanna?

The Hierarchical Patch Dynamics Paradigm provides a conceptual framework for linking pattern, process and scale in ecosystems, but there have been few attempts to test this theory because most ecological studies focus on only one spatial scale, or are limited in their temporal scope. Here I use palaeoecological techniques (analysis of fossil pollen and stable carbon isotopes) to compare vegetation heterogeneity in an east African savanna at three spatial scales, over hundreds of years. The data show that patterns of vegetation change are different at the three spatial scales of observation, and suggest that different ecological processes dominate tree abundance at micro, local and landscape scales. Interactions between plants, disturbance (e.g., by fire and herbivores), climate and soil type may influence tree density at differing spatial and temporal scales. This hierarchical explanation of savanna vegetation dynamics could inform future biodiversity conservation and management in savannas. This revised version was published online in July 2006 with corrections to the Cover Date.     

Modeling scale-dependent landscape pattern,dispersal, and connectivity from the perspective of the organism

Understanding the impacts of habitat fragmentation on dispersal is an important issue in landscape and conservation ecology. Here I examine the effects of fine- to broad-scale patterns in landscape structure on dispersal success of organisms with differing life-history traits. An individual-based model was used to simulate dispersal of amphibian-like species whose movements were driven by land cover and moisture conditions. To systematically control spatial pattern, a landscape model was created by merging simulated land cover maps with synthetic topographic surfaces. Landscapes varied in topographic roughness and spatial contagion in agriculture and urban land cover. Simulations included three different species types that varied in their maximum potential dispersal distances by 1-, 2-, or 4-fold. Two sets of simulations addressed effects of varying aspects of landscape structure on dispersal success. In the first set of simulations, which incorporated variable distances between breeding patches, dispersal success was lowest for all species types when anthropogenic cover was patchily distributed. In the second set, with interpatch distances held constant as landscape composition varied, dispersal success decreased as anthropogenic cover became spatially contagious. Both sets revealed strong main effects of species characteristics, interpatch distances and landscape composition on dispersal success; furthermore, scale-dependent patterns in land cover and moisture gradients had a stronger effect on longer- than shorter-ranging species types. Taken together, these simulations suggest that heuristic conservation strategies could potentially be developed based on important but limited life history information.     

Landscape effects on scales of movement by white-tailed deer in an agricultural–forest matrix

Understanding how organisms respond to landscape heterogeneity is foundational to landscape ecology. We characterized seasonal scales of movement of white-tailed deer (Odocoileus viginianus) in an agricultural–forest matrix using first-passage time analysis (FPT) for 62 GPS-collared individuals. We investigated whether those scales were driven by demographic or landscape features. We found FPT for each individual across all seasons was typically dominated by a peak in variance of FPT/area at scales (radii) from 425 to 1,675 m. These peaks occurred at scales consistent with seasonal space use. We observed additional lower magnitude peaks at larger scales (3,000–6,000 m) and small scales (25–150 m). Peaks at larger scales were associated with seasonal migrations and dispersal events. Small scale peaks may represent resting or foraging behavior. Female movements were organized at smaller scales than males in the spring/summer season. Models relating landscape features to movement scales suggest that deer perceive and move within the landscape differently as the roles of dominant land-cover types shift seasonally. During winter, configuration (interspersion/juxtaposition) of land-cover types is more important to deer than during spring/summer and fall. During spring/summer and fall, movement behavior may be dictated by reproductive and harvest activities.     

Dispersion of kangaroo rat mounds at multiple scales in New Mexico,USA

Burrowing mammals create disturbances that increase the ecological heterogeneity of landscapes. In desert systems, banner-tailed kangaroo rats (Dipodomys spectabilis) construct large mounds that greatly influence the spatial patterning of soils, plants, and animals. The overall effects of the patches generated by D. spectabilis depend on the dispersion patterns of the mounds; these patterns may be sensitive to scale and landscape position. We examined the distribution of D. spectabilis mounds across multiple scales in four 40-ha grassland plots in New Mexico, USA. We used Ripley's K-function for our point-pattern analysis. The dispersion patterns of mounds were generally scale-sensitive but depended somewhat on plot-level densities, which were related to topographic position and grazing history. Mound spacing was either regular or random at small scales (0–50 m), random or aggregated at intermediate scales (50–300 m), and aggregated at large scales (300–3000 m). This scale-dependency of pattern reflected spatial domains in which different biotic (territoriality, dispersal, grazing) and abiotic (soil texture and drainage) factors exerted strong influences. How other organisms perceive the spatial patterning of mounds will depend on the extent of their movements. Patches may appear regular to one species but aggregated to another. The dispersion of D. spectabilis mounds also has implications for the spatial structuring of desert rodent communities. D. spectabilis excludes smaller species of kangaroo rats from areas around their mounds; they create spatial heterogeneity in behavioral dominance that may influence the distribution of subordinate species at multiple scales.     

Foraging in a fractal environment: Spatial patterns in a marine predator-prey system

Spatial relationships between predators and prey have important implications for landscape processes and patterns. Highly mobile oceanic birds and their patchily distributed prey constitute an accessible model system for studying these relationships. High-frequency echosounders can be used together with simultaneous direct visual observations to quantitatively describe the distributions of seabird consumers and their resources over a wide range of spatial scales, yielding information which is rarely available in terrestrial systems.Recent fine-scale investigations which have used acoustics to study the distribution of foraging marine birds have reported weak or ephemeral spatial associations between the birds and their prey. These results are inconsistent with predictions of optimal foraging, but several considerations suggest that traditional foraging models do not adequately describe resource acquisition in marine environments. Relative to their terrestrial counterparts, oceanic landscapes are structurally very simple, but they generally lack visual cues about resource availability.An emerging view assumes that perceptually constrained organisms searching for food in multiscale environments should respond to patterns of resource abundance over a continuum of scales. We explore fractal geometry as a possible tool for quantifying this view and for describing spatial dispersion patterns that result from foraging behavior. Data on an Alaskan seabird (least auklet [Aethia pusilla]) and its zooplanktonic food resources suggest that fractal approaches can yield new ecological insights into complex spatial patterns deriving from animal movements.     

Topographic placement of management practices in riparian zones to reduce water quality impacts from pastures

Riparian buffers, a best management practice (BMP), lessen environmental impacts caused by pasture-based agriculture by excluding livestock from streams and removing sediment and nutrients from overland flow before they enter the watercourse. Geospatial data analysis of digital elevation models (DEMs) can improve BMP placement by ensuring that BMPs intercept flow paths. Livestock heavy use areas and riparian buffers within 100?m of the stream were digitized using aerial photography of Spring Creek watershed (Pennsylvania, USA). Flow path lengths and stream entry points from heavy use areas, from all agricultural land uses, and from a random sample of points were calculated using 1, 10, and 30?m DEM resolutions. Euclidean distances identified the nearest stream entry point regardless of topography. Drainage areas for each streambank cell were evaluated using each DEM resolution. Topographic calculations differed significantly from Euclidean, with median differences of 14.3?m for flow path length and 24.1?m between stream entry points for the 10?m DEM. Existing buffers intercepted runoff from only 23?% of heavy use areas. Drainage areas ranged from one to hundreds of DEM cells. Any DEM is an arbitrary representation of a continuous surface; both resolution and processing method affect the suitability of such a representation for any given purpose. We found that 30?m DEMs did not provide reasonable flow path estimates at the scale of grazing agriculture in this region, while Pennsylvania 1?m DEMs were minimally smoothed during DEM preparation, resulting in erratic flow paths. The 10?m DEMs were the most suitable available DEM product, and should be used in conjunction with site visits for planning pasture BMP placement.     

Using network theory to prioritize management in a desert bighorn sheep metapopulation

Connectivity models using empirically-derived landscape resistance maps can predict potential linkages among fragmented animal and plant populations. However, such models have rarely been used to guide systematic decision-making, such as identifying the most important habitat patches and dispersal corridors to protect or restore in order to maximize regional connectivity. Combining resistance models with network theory offers one means of prioritizing management for connectivity, and we applied this approach to a metapopulation of desert bighorn sheep (Ovis canadensis nelsoni) in the Mojave Desert of the southwestern United States. We used a genetic-based landscape resistance model to construct network models of genetic connectivity (potential for gene flow) and demographic connectivity (potential for colonization of empty habitat patches), which may differ because of sex-biased dispersal in bighorn sheep. We identified high-priority habitat patches and corridors and found that the type of connectivity and the network metric used to quantify connectivity had substantial effects on prioritization results, although some features ranked highly across all combinations. Rankings were also sensitive to our empirically-derived estimates of maximum effective dispersal distance, highlighting the importance of this often-ignored parameter. Patch-based analogs of our network metrics predicted both neutral and mitochondrial genetic diversity of 25 populations within the study area. This study demonstrates that network theory can enhance the utility of landscape resistance models as tools for conservation, but it is critical to consider the implications of sex-biased dispersal, the biological relevance of network metrics, and the uncertainty associated with dispersal range and behavior when using this approach.     

Scales of movement by elk (<Emphasis Type="Italic">Cervus elaphus</Emphasis>) in response to heterogeneity in forage resources and predation risk

Animals may respond to spatial and temporal heterogeneity by altering their movement patterns. The time an animal spends in an area of a given size is termed ȁ8first-passage timeȁ9 and can be used to identify the scales at which different movement processes occur. Using first-passage time and 2-h observations, we identified nested spatial scales representing three movement behaviours for elk (Cervus elaphus) – inactive/resting (moves < 50 m), active/foraging (x̄ = 276.7 m, SD = 56.6), and active/relocating (x̄ = 1628.3 m, SD = 436.6). Our ability to identify inactive behaviour was limited by GPS accuracy. The scale separating relocating and foraging behaviour ranged 550–1650 m across individuals and varied quadratically with the mean patch size of cutover forest in an animal’s home range. We classified path segments into the 3 movement behaviours and related behaviours to local environmental conditions. Elk were likely to be inactive in areas having a low predicted use by wolves (Canis lupus), farther than 50 m from anthropogenic linear clearings, and where microclimatic conditions were cool (high shrub cover and north to east-facing slopes). In contrast, elk were most likely to forage in areas having intermediate levels of herbaceous biomass and low movement costs. Elk were most likely to be relocating when in areas of high wolf use, when close to linear clearings, and in energetically costly situations such as moving upslope. We discuss how elk use of potential foraging habitats may be restricted in this landscape by risks imposed by predators, humans, or both.