Hierarchical habitat selection by woodland caribou: its relationship to limiting factors

Habitat selection is a hierarchical process that may yield various patterns depending on the scales of investigation. We employed satellite radio‐telemetry to examine patterns of habitat selection by female woodland caribou in central Saskatchewan at both coarse (seasonal range) and fine (daily area) scales. At each scale, we converted spatial data describing compositions of available and used habitat to standardised resource selection indices and examined them with multivariate analyses of variance. Seasonal ranges generally showed preferential inclusion of peatlands and black spruce dominated stands relative to recently disturbed stands and early seral stage forests. In all populations, caribou preferred peatlands and black spruce forests to all other habitat types at the daily area scale, in general, these patterns may reveal the effective avoidance of wolves, the primary factor limiting caribou throughout the boreal forest. In three populations where seasonal ranges showed the selective inclusion of either young jack pine stands or clearcuts along with peatlands and black spruce forests, we found a relative avoidance of the clearcuts and young jack pine stands at the daily area scale. As all caribou populations in the area are thought to be relics of a once more continuous distribution, the seasonal range selection by animals in disturbed areas may better describe historic rather than current habitat selection. We found inter‐annual variation in selection at the coarser spatial scale in one population, and inter‐seasonal variation in selection at the finer spatial scale in three populations, indicating that the relative grains of the spatial and temporal scales coincide. We were better able to explain the seasonal variations in finer scale selection by considering available forage, a factor less likely than predation to limit woodland caribou populations. The data agree with the theory that the spatial and temporal hierarchy of habitat selection reflects the hierarchy of factors potentially limiting individual fitness.     

Scale for resource selection functions

Resource selection functions (RSFs) are statistical models defined to be proportional to the probability of use of a resource unit. My objective with this review is to identify how RSFs can be used to unravel the influence of scale in habitat selection. In wildlife habitat studies, including radiotelemetry, RSFs can be estimated using a variety of statistical methods, all of which can be used to explore the role of scale. All RSFs are bounded by the resolution of data and the spatial extent of the study area, but also allow predictor covariates to be measured at a variety of scales. Conditional logistic regression permits designs (e.g. matched case) that relate the process of habitat selection to a limited domain of resource units that might better characterize what is truly ‘available’ to the animal. Scale influences the process of habitat selection, e.g. food resources are often selected at fine spatial scales, whereas landscape patterns at much larger scales typically influence the location of home ranges. Scale also influences appropriate sampling in many ways: (1) heterogeneity might be obliterated (transmutation) if resolution or grain size is too large, (2) variance of habitat characteristics might be undersampled if extent or domain is too small, (3) timing and duration of observations can influence RSF models, and (d) both spatial and temporal autocorrelations can vary directly with the intensity of sampling. Using RSFs, researchers can examine habitat selection at multiple scales, and predictive models that bridge scales can be estimated. Using Geographical Information Systems, predictor covariates in RSF models can be measured at different scales easily so that the predictive ability of models at alternative spatial and temporal domains can be explored by the investigator. Identification of the scale that best explains the data can be evaluated by comparing alternative models using information‐theoretic metrics such as Akaike Information Criteria, and predictive capability of the models can be assessed using k‐fold cross validation.     

Selecting Habitat to Survive: The Impact of Road Density on Survival in a Large Carnivore

Habitat selection studies generally assume that animals select habitat and food resources at multiple scales to maximise their fitness. However, animals sometimes prefer habitats of apparently low quality, especially when considering the costs associated with spatially heterogeneous human disturbance. We used spatial variation in human disturbance, and its consequences on lynx survival, a direct fitness component, to test the Hierarchical Habitat Selection hypothesis from a population of Eurasian lynx Lynx lynx in southern Norway. Data from 46 lynx monitored with telemetry indicated that a high proportion of forest strongly reduced the risk of mortality from legal hunting at the home range scale, while increasing road density strongly increased such risk at the finer scale within the home range. We found hierarchical effects of the impact of human disturbance, with a higher road density at a large scale reinforcing its negative impact at a fine scale. Conversely, we demonstrated that lynx shifted their habitat selection to avoid areas with the highest road densities within their home ranges, thus supporting a compensatory mechanism at fine scale enabling lynx to mitigate the impact of large-scale disturbance. Human impact, positively associated with high road accessibility, was thus a stronger driver of lynx space use at a finer scale, with home range characteristics nevertheless constraining habitat selection. Our study demonstrates the truly hierarchical nature of habitat selection, which aims at maximising fitness by selecting against limiting factors at multiple spatial scales, and indicates that scale-specific heterogeneity of the environment is driving individual spatial behaviour, by means of trade-offs across spatial scales.     

An Examination of Scale-Dependent Resource Use by Eastern Hognose Snakes in Southcentral New Hampshire

ABSTRACT The decline of many snake populations is attributable to habitat loss, and knowledge of habitat use is critical to their conservation. Resource characteristics (e.g., relative availability of different habitat types, soils, and slopes) within a landscape are scale-dependent and may not be equal across multiple spatial scales. Thus, it is important to identify the relevant spatial scales at which resource selection occurs. We conducted a radiotelemetry study of eastern hognose snake (Heterodon platirhinos) home range size and resource use at different hierarchical spatial scales. We present the results for 8 snakes radiotracked during a 2-year study at New Boston Air Force Station (NBAFS) in southern New Hampshire, USA, where the species is listed by the state as endangered. Mean home range size (minimum convex polygon) at NBAFS (51.7 ± 14.7 ha) was similar to that reported in other parts of the species’ range. Radiotracked snakes exhibited different patterns of resource use at different spatial scales. At the landscape scale (selection of locations within the landscape), snakes overutilized old-field and forest edge habitats and underutilized forested habitats and wetlands relative to availability. At this scale, snakes also overutilized areas containing sandy loam soils and areas with lower slope (mean slope = 5.2% at snake locations vs. 6.7% at random locations). We failed to detect some of these patterns of resource use at the home range scale (i.e., within the home range). Our ability to detect resource selection by the snakes only at the landscape scale is likely the result of greater heterogeneity in macrohabitat features at the broader landscape scale. From a management perspective, future studies of habitat selection for rare species should include measurement of available habitat at spatial scales larger than the home range. We suggest that the maintenance of open early successional habitats as a component of forested landscapes will be critical for the persistence of eastern hognose snake populations in the northeastern United States.     

Using multilevel models to quantify heterogeneity in resource selection

Models of resource selection are being used increasingly to predict or model the effects of management actions rather than simply quantifying habitat selection. Multilevel, or hierarchical, models are an increasingly popular method to analyze animal resource selection because they impose a relatively weak stochastic constraint to model heterogeneity in habitat use and also account for unequal sample sizes among individuals. However, few studies have used multilevel models to model coefficients as a function of predictors that may influence habitat use at different scales or quantify differences in resource selection among groups. We used an example with white-tailed deer (Odocoileus virginianus) to illustrate how to model resource use as a function of distance to road that varies among deer by road density at the home range scale. We found that deer avoidance of roads decreased as road density increased. Also, we used multilevel models with sika deer (Cervus nippon) and white-tailed deer to examine whether resource selection differed between species. We failed to detect differences in resource use between these two species and showed how information-theoretic and graphical measures can be used to assess how resource use may have differed. Multilevel models can improve our understanding of how resource selection varies among individuals and provides an objective, quantifiable approach to assess differences or changes in resource selection. © 2011 The Wildlife Society.     

Scale-dependent selection of greenness by African elephants in the Kruger-private reserve transboundary region,South Africa

Foraging behaviour and habitat selection occur as hierarchical processes. Understanding the factors that govern foraging and habitat selection thus requires investigation of those processes over the scales at which they occur. We investigated patterns of habitat use by African elephants (Loxodonta africana) in relation to vegetation greenness to investigate the scale at which that landscape attribute was most closely related to distribution of elephant locations. We analysed Global Positioning System radio-collar locations for 15 individuals, using the Normalized Difference Vegetation Index as a representation of vegetation greenness in a Geographic Information Systems framework. We compared the importance of vegetation greenness at three spatial scales: the total home range, the seasonal home range and the 16-day home range. During the wet season, seasonal home ranges for both sexes were associated with intermediate greenness within the total home range; there was no evidence of selection based on greenness at finer scales. During the dry season, the strongest associations were within the 16-day home range: individual locations for males tended to be in areas of intermediate greenness, and those for females were in areas of intermediate and high greenness. Our findings suggest that the role of vegetation greenness varies with the scale of analysis, likely reflecting the hierarchical processes involved in habitat selection by elephants.     

Linking moose habitat selection to limiting factors

It has been suggested that patterns of habitat selection of animals across spatial scales should reflect the factors limiting individual fitness in a hierarchical fashion. Animals should thus select habitats that permit avoidance of the most important limiting factor at large spatial scales while the influence of less important factors should only be evident at fine scales. We tested this hypothesis by investigating moose Alces alces habitat selection using GPS telemetry in an area where the main factors limiting moose numbers were likely (in order of decreasing importance) predation risk, food availability and snow. At the landscape scale, we predicted that moose would prefer areas where the likelihood of encountering wolves was low or areas where habitats providing protection from predation were dominant. At the home‐range scale, we predicted that moose selection would be driven by food availability and snow depth. Wolf territories were delineated using telemetry locations and the study area was divided into 3 sectors that differed in terms of annual snowfall. Vegetation surveys yielded 6 habitat categories that differed with respect to food availability, and shelter from predation or snow. Our results broadly supported the hypothesis because moose reacted to several factors at each scale. At the landscape scale, moose were spatially segregated from wolves by avoiding areas receiving the lowest snowfall, but they also preferentially established their home range in areas where shelter from snow bordered habitat types providing abundant food. At the home‐range scale, moose also traded off food availability with avoidance of deep snow and predation risk. During winter, moose increased use of stands providing shelter from snow along edges with stands providing abundant food. Habitat selection patterns of females with calves differed from that of solitary moose, the former being associated primarily with habitats providing protection from predation. Animals should attempt to minimize detrimental effects of the main limiting factors when possible at the large scale. However, when the risk associated with several potential limiting factors varies with scale, we should expect animals to make trade‐offs among these.     

Scales and costs of habitat selection in heterogeneous landscapes

Summary Two scales of habitat selection are likely to influence patterns of animal density in heterogeneous landscapes. At one scale, habitat selection is determined by the differential use of foraging locations within a home range. At a larger scale, habitat selection is determined by dispersal and the ability to relocate the home range. The limits of both scales must be known for accurate assessments of habitat selection and its role in effecting spatial patterns in abundance. Isodars, which specify the relationships between population density in two habitats such that the expected reproductive success of an individual is the same in both, allow us to distinguish the two scales of habitat selection because each scale has different costs. In a two-habitat environment, the cost of rejecting one of the habitats within a home range can be expressed as a devaluation of the other, because, for example, fine-grained foragers must travel through both. At the dispersal scale, the cost of accepting a new home range in a different habitat has the opposite effect of inflating the value of the original habitat to compensate for lost evolutionary potential associated with relocating the home range. These costs produce isodars at the foraging scale with a lower intercept and slope than those at the dispersal scale.Empirical data on deer mice occupying prairie and badland habitats in southern Alberta confirm the ability of isodar analysis to differentiate between foraging and dispersal scales. The data suggest a foraging range of approximately 60 m, and an effective dispersal distance near 140 m. The relatively short dispersal distance implies that recent theories may have over-emphasized the role of habitat selection on local population dynamics. But the exchange of individuals between habitats sharing irregular borders may be substantial. Dispersal distance may thus give a false impression of the inability of habitat selection to help regulate population density.     

Wolf pack rendezvous site selection in Greece is mainly affected by anthropogenic landscape features

In wolves, most offspring mortality occurs within the first 6–8 months of their life. As wolf pups pass this entire period at either the den or rendezvous sites, their selection by wolf packs may affect pup survival and recruitment. Rendezvous sites are important for pup survival as they are used during summer and early autumn, when intense human activity may increase pup mortality. Adult wolves and pups can be killed by livestock guarding dogs during summer and intentionally or accidentally during large game hunting in autumn. This study describes factors related to rendezvous site selection in order to enhance their protection and management. We studied the rendezvous site selection of 30 wolf packs in central and northern Greece between 1998 and 2010, after locating 35 sites using the simulated howling survey method and telemetry. We considered a series of environmental and anthropogenic predictors of wolf rendezvous site selection at two spatial scales. At the landscape-population scale, wolves selected rendezvous sites below 1,200 m asl, with large inter-site distance (mean, 12.9 km), and avoided partially forested or open habitats, indicating preference for covered, spaced areas with seasonally stable resources. At the home range scale, wolves selected rendezvous sites away from forest roads and villages, close to water sources, and in areas with low forest fragmentation, indicating avoidance of human presence and disturbance. In the summer of 2011, we used an ensuing resource selection model (RSF, AUC?=?0.818) to successfully locate seven new rendezvous sites outside our previous survey area, verifying the utility of prediction maps (all new sites were at areas with 0.8–1 model probability). Rendezvous prediction maps can be used to reduce field effort when monitoring wolf populations, assess livestock predation risk, design protected areas, and reduce human disturbance on reproductive wolf packs.     

Scale dependency and functional response in moose habitat selection

Habitat selection can be influenced by the distribution of the habitat types in the landscape as well as net gain in visiting patches of resources, causing individual variation in habitat selection. Moreover, the hypothesis of functional response in habitat selection predicts that the degree of selection of a resource depends on its relative availability. We used radio-telemetry data from individual moose on an island off the coast of northern Norway to evaluate whether the selection of habitat types at the landscape scale differed from the choice of habitat types within the home range, and investigated the functional response in habitat selection by relating individual habitat selection to home range characteristics. At the landscape scale, moose selected for habitat types that provided both good forage and cover, with small differences between sex and age groups. At the home range scale, all individuals selected habitat types that were associated with cover and low human impact. Habitat selection was not modified by local moose density, but was related to home range size at both spatial scales. Larger home ranges contained larger proportions of non-preferred habitat types compared to smaller home ranges. At the home range scale, the selection for a habitat type decreased with its relative availability, indicating a functional response in habitat selection. This suggests that habitat selection is modified by home range size, which influences the availability of habitat types and shapes individual habitat selection patterns. Our results support previous suggestions that analyses of habitat or resource selection should follow a multi-scale approach. Both the relative availability of habitat types as well as individual variation in home range size should be accounted for in order to disentangle the complex mechanisms that contribute to shape patterns of resource selection in animals.     

Habitat selection by a large herbivore at multiple spatial and temporal scales is primarily governed by food resources

Habitat selection can be considered as a hierarchical process in which animals satisfy their habitat requirements at different ecological scales. Theory predicts that spatial and temporal scales should co‐vary in most ecological processes and that the most limiting factors should drive habitat selection at coarse ecological scales, but be less influential at finer scales. Using detailed location data on roe deer Capreolus capreolus inhabiting the Bavarian Forest National Park, Germany, we investigated habitat selection at several spatial and temporal scales. We tested 1) whether time‐varying patterns were governed by factors reported as having the largest effects on fitness, 2) whether the trade‐off between forage and predation risks differed among spatial and temporal scales and 3) if spatial and temporal scales are positively associated. We analysed the variation in habitat selection within the landscape and within home ranges at monthly intervals, with respect to land‐cover type and proxys of food and cover over seasonal and diurnal temporal scales. The fine‐scale temporal variation follows a nycthemeral cycle linked to diurnal variation in human disturbance. The large‐scale variation matches seasonal plant phenology, suggesting food resources being a greater limiting factor than lynx predation risk. The trade‐off between selection for food and cover was similar on seasonal and diurnal scale. Habitat selection at the different scales may be the consequence of the temporal variation and predictability of the limiting factors as much as its association with fitness. The landscape of fear might have less importance at the studied scale of habitat selection than generally accepted because of the predator hunting strategy. Finally, seasonal variation in habitat selection was similar at the large and small spatial scales, which may arise because of the marked philopatry of roe deer. The difference is supposed to be greater for wider ranging herbivores.     

A Hierarchical Analysis of Habitat Selection by Raccoons in Northern Indiana

Abstract: Although numerous studies have examined habitat use by raccoons (Procyon lotor), information regarding seasonal habitat selection related to resource availability in agricultural landscapes is lacking for this species. Additionally, few studies using radiotelemetry have investigated habitat selection at multiple spatial scales or core-use areas by raccoons. We examined seasonal habitat selection of 55 (31 M, 24 F) adult raccoons at 3 hierarchical orders defined by the movement behavior of this species (second-order home range, second-order core-use area, and third-order home range) in northern Indiana, USA, from May 2003 to June 2005. Using compositional analysis, we assessed whether habitat selection differed from random and ranked habitat types in order of selection during the crop growing period (season 1) and corn maturation period (season 2), which represented substantial shifts in resource availability to raccoons. Habitat rankings differed across hierarchical orders, between seasons within hierarchical orders, and between sexes within seasons; however, seasonal and intersexual patterns of habitat selection were not consistent across hierarchical orders of spatial scale. When nonrandom utilization was detected, both sexes consistently selected forest cover over other available habitats. Seasonal differences in habitat selection were most evident at the core-area scale, where raccoon selection of agricultural lands was highest during the maturation season when corn was available as a direct food source. Habitat use did not differ from availability for either sex in either season at the third-order scale. The selection of forest cover across both seasons and all spatial orders suggested that raccoon distribution and abundance in fragmented landscapes is likely dependent on the availability and distribution of forest cover, or habitats associated with forest (i.e., water), within the landscape. The lack of consistency in habitat selection across hierarchical scales further exemplifies the need to examine multiple biological scales in habitat-selection studies.     

Home range and habitat selection of spotted owls in the central Sierra Nevada

We studied home range and habitat selection of radio-marked adult California spotted owls (Strix occidentalis occidentalis) randomly selected from among the breeding population of owls in the central Sierra Nevada, California from June to October 2006. The most parsimonious home-range estimate for our data was 555 ha (SE = 100 ha). Home-range size was positively correlated with the number of vegetation patches in the home range (habitat heterogeneity). We used resource selection ratios to examine selection of vegetation types by owls within our study area. Owl home ranges contained a high proportion of mature conifer forest, relative to its availability, although the confidence interval for this estimate overlapped one. We also used resource selection functions (RSF) to examine owl foraging habitat selection. Relative probability of selection of foraging habitat was correlated with vegetation classes, patch size, and their interaction. Owls showed highest selection rates for large patches (>10 ha) of pole-sized coniferous forest. Our results suggested that spotted owls in the central Sierra Nevada used habitat that contained a high proportion of mature conifer forest at the home-range scale, but at a finer scale (foraging site selection) owls used other vegetation classes interspersed among mature forest patches, consistent with our hypothesis that spotted owls may use other forest types besides old growth and mature forests when foraging. Our study provides an unbiased estimate of habitat use by spotted owls in the central Sierra Nevada. Our results suggest that forest managers continue to protect remaining mature and old-growth forests in the central Sierra Nevada because owl home ranges contain high proportions of these habitats. However, our results also showed that owls used younger stands as foraging habitat so that landscape heterogeneity, with respect to cover types, may be an important consideration for management but we did not attempt to relate our findings to fitness of owls. Thus management for some level of landscape heterogeneity for the benefit of owls should proceed with caution or under an adaptive management framework. © 2011 The Wildlife Society.     

Habitat selection as a hierarchy: the spatial scales of winter foraging by muskoxen

We studied the winter resource selection of muskoxen Ovibos moschatus in the High Arctic using a nested hierarchy of spatial scales 1) population range, 2) travel routes, 3) feeding sites (l e clusters of feeding craters), 4) feeding craters, and 5) diet (I e plant species) We found that, generally, patterns of selection remained consistent across all levels At successively smaller scales, muskoxen selected for higher graminoid abundance and particularly for thinner, softer snow cover, although we did not reject the hypothesis of random travel route selection Muskoxen uncovered forages from beneath the snow cover, by cratering, near the flonstic and nival extremes of availability Selection was consistently biased toward use of water sedge, Carex aquatilis As scale changed, however, muskoxen showed reversals of preference for some other forage species Diet was dominated by C aquatilis and cotton sedge, Eriophorum angustifolium , species characteristic of lowland meadows During spring melt, muskoxen moved to snow-free uplands to feed Dietary quality, as revealed by fecal nitrogen, increased at this time The consistency of the results across scales implied that these local levels of habitat selection occurred within one scaling domain     

Habitat Selection by American Martens in Coastal California

ABSTRACT We investigated habitat selection using single- and mixed-scale modeling at 2 spatial scales, stand and home range, by the only known population of American martens (Martes americana) remaining in the historical range of the Humboldt subspecies (M. a. humboldtensis) in California, USA. During 2000 and 2001, we sampled a 12 times 14 grid with 2-km spacing, using 2 sooted track plates at each grid point. We detected martens at 26 of the 159 grid points. We used resource selection probability functions and an information-theoretic method to model habitat at detection locations. At the stand scale, martens selected conifer-dominated stands with dense, spatially extensive shrub cover (x̄ = 74% cover, SE = 4) in the oldest developmental stage. At the home-range scale, martens selected the largest available patches (x̄ = 181 ha, SE = 14) of old-growth, old-growth and late-mature, or serpentine habitat. Mixed-scale models revealed that habitat characteristics from both scales best explained marten occurrence compared to one scale alone. Dense, spatially extensive shrub cover is a key habitat element for martens in coastal forests. Dense shrubs provide refuge from predators, cover for prey, and may also deter larger-bodied competitors. Managers can increase the likelihood of marten population persistence and encourage expansion in coastal forests by maintaining and restoring late-mature and old-growth, conifer-dominated forests with dense shrub cover in large, contiguous patches.     

The ecology of 3-D space use in a sexually dimorphic mammal

The distribution of animals is the result of habitat selection according to sex, reproductive status and resource availability. Little is known about how marine predators investigate their 3-dimensional space along both the horizontal and vertical axes and how temporal variation affects space use. In this study, we assessed the spatio-temporal movement of a sexually dimorphic marine mammal, the grey seal Halichoerus grypus by 1) determining seasonal home range size, 2) testing whether space use of seals was affected by water depth, and 3) investigating the vertical movement of seals according to the maximum depth of each dive. Between 1993 and 2005, we fitted 49 grey seals in the Gulf of St. Lawrence with satellite transmitters. We estimated seasonal 95% fixed-kernel home ranges for each individual. For each seal, we tested for selectivity and preference for 4 water depth classes at the home range scale and within the home range. We also evaluated the proportional number of dives made in each water depth classes according to the maximum depth of each dive. Home ranges were 10 times larger in winter than in summer. Seals generally selected habitats <50 m deep. They also mainly dove to depths of 40 m or less. At both scales of selection, preference for shallow areas decreased in winter. We also observed that adults used shallow habitats more than juveniles to establish their home range. A spatial segregation based on sex also occurred at the finer scale of selection where females were more concentrated in the shallowest parts of their home range than males. Segregation in space use according to age and sex classes occurred at both the horizontal and vertical scales. Our results emphasise the importance of studying habitat selection of marine predators in 3-dimensional space, in addition to the temporal scale.     

Correlation and studies of habitat selection: problem,red herring or opportunity?

With the advent of new technologies, animal locations are being collected at ever finer spatio-temporal scales. We review analytical methods for dealing with correlated data in the context of resource selection, including post hoc variance inflation techniques, ‘two-stage’ approaches based on models fit to each individual, generalized estimating equations and hierarchical mixed-effects models. These methods are applicable to a wide range of correlated data problems, but can be difficult to apply and remain especially challenging for use–availability sampling designs because the correlation structure for combinations of used and available points are not likely to follow common parametric forms. We also review emerging approaches to studying habitat selection that use fine-scale temporal data to arrive at biologically based definitions of available habitat, while naturally accounting for autocorrelation by modelling animal movement between telemetry locations. Sophisticated analyses that explicitly model correlation rather than consider it a nuisance, like mixed effects and state-space models, offer potentially novel insights into the process of resource selection, but additional work is needed to make them more generally applicable to large datasets based on the use–availability designs. Until then, variance inflation techniques and two-stage approaches should offer pragmatic and flexible approaches to modelling correlated data.     

The scale-dependent impact of wolf predation risk on resource selection by three sympatric ungulates

Resource selection is a fundamental ecological process impacting population dynamics and ecosystem structure. Understanding which factors drive selection is vital for effective species- and landscape-level management. We used resource selection probability functions (RSPFs) to study the influence of two forms of wolf (Canis lupus) predation risk, snow conditions and habitat variables on white-tailed deer (Odocoileus virginianus), elk (Cervus elaphus) and moose (Alces alces) resource selection in central Ontario's mixed forest French River-Burwash ecosystem. Direct predation risk was defined as the frequency of a predator's occurrence across the landscape and indirect predation risk as landscape features associated with a higher risk of predation. Models were developed for two winters, each at two spatial scales, using a combination of GIS-derived and ground-measured data. Ungulate presence was determined from snow track transects in 64 16- and 128 1-km(2) resource units, and direct predation risk from GPS radio collar locations of four adjacent wolf packs. Ungulates did not select resources based on the avoidance of areas of direct predation risk at any scale, and instead exhibited selection patterns that tradeoff predation risk minimization with forage and/or mobility requirements. Elk did not avoid indirect predation risk, while both deer and moose exhibited inconsistent responses to this risk. Direct predation risk was more important to models than indirect predation risk but overall, abiotic topographical factors were most influential. These results indicate that wolf predation risk does not limit ungulate habitat use at the scales investigated and that responses to spatial sources of predation risk are complex, incorporating a variety of anti-predator behaviours. Moose resource selection was influenced less by snow conditions than cover type, particularly selection for dense forest, whereas deer showed the opposite pattern. Temporal and spatial scale influenced resource selection by all ungulate species, underlining the importance of incorporating scale into resource selection studies.     

At what spatial scales does resource selection vary? A case study of koalas in a semi‐arid region

The conservation of any species requires understanding and predicting the distribution of its habitat and resource use, including the effects of scale‐dependent variation in habitat and resource quality. Consequently, testing for resource selection at the appropriate scales is critical. We investigated how the resource selection process varies across scales, using koalas in a semi‐arid landscape of eastern Australia as a case study. We asked: at what scales does tree selection by koalas vary across regions? We tested the importance of the variation of our ecological predictors at the following scales: (i) the site‐scale (a stand of trees representing an individual koala's perception of local habitat); (ii) the landscape‐scale (10 × 10 km area representing a space within which a population of koalas exists); and (iii) a combination of these scales. We used a mixed‐modelling approach to quantify variation in selection of individual trees by koalas among sites and landscapes within a 1600 km² study area. We found that tree species, and tree height, were the most important factors influencing tree selection, and that their effect did not vary across scales. In contrast, preferences for trees of different condition, which is the state of tree canopy health, did vary across landscapes, indicating spatial variation in the selection of trees with respect to tree condition at the landscape‐scale, but not at the site‐scale. We conclude that resource selection processes can depend on the quality of those resources at different scales and their heterogeneous nature across landscapes, highlighting the consequence of scale‐dependent ecological processes. Designing studies that capture the heterogeneity in habitat and resources used by species that have an extensive distribution is an important prerequisite for effective conservation planning and management.     

Scale Dependence of Female Ungulate Reproductive Success in Relation to Nutritional Condition,Resource Selection and Multi-Predator Avoidance

Female ungulate reproductive success is dependent on the survival of their young, and affected by maternal resource selection, predator avoidance, and nutritional condition. However, potential hierarchical effects of these factors on reproductive success are largely unknown, especially in multi-predator landscapes. We expanded on previous research of neonatal white-tailed deer (Odocoileus virginianus) daily survival within home ranges to assess if resource use, integrated risk of 4 mammalian predators, maternal nutrition, winter severity, hiding cover, or interactions among these variables best explained landscape scale variation in daily or seasonal survival during the post-partum period. We hypothesized that reproductive success would be limited greater by predation risk at coarser spatiotemporal scales, but habitat use at finer scales. An additive model of daily non-ideal resource use and maternal nutrition explained the most (69%) variation in survival; though 65% of this variation was related to maternal nutrition. Strong support of maternal nutrition across spatiotemporal scales did not fully support our hypothesis, but suggested reproductive success was related to dam behaviors directed at increasing nutritional condition. These behaviors were especially important following severe winters, when dams produced smaller fawns with less probability of survival. To increase nutritional condition and decrease wolf (Canis lupus) predation risk, dams appeared to place fawns in isolated deciduous forest patches near roads. However, this resource selection represented non-ideal resources for fawns, which had greater predation risk that led to additive mortalities beyond those related to resources alone. Although the reproductive strategy of dams resulted in greater predation of fawns from alternative predators, it likely improved the life-long reproductive success of dams, as many were late-aged (>10 years old) and could have produced multiple litters of fawns. Our study emphasizes understanding the scale-dependent hierarchy of factors limiting reproductive success is essential to providing reliable knowledge for ungulate management.