Living on the edge: Multiscale habitat selection by cheetahs in a human‐wildlife landscape

Animals select habitats that will ultimately optimize their fitness through access to favorable resources, such as food, mates, and breeding sites. However, access to these resources may be limited by bottom‐up effects, such as availability, and top‐down effects, such as risk avoidance and competition, including that with humans. Competition between wildlife and people over resources, specifically over space, has played a significant role in the worldwide decrease in large carnivores. The goal of this study was to determine the habitat selection of cheetahs (Acinonyx jubatus) in a human‐wildlife landscape at multiple spatial scales. Cheetahs are a wide‐ranging, large carnivore, whose significant decline is largely attributed to habitat loss and fragmentation. It is believed that 77% of the global cheetah population ranges outside protected areas, yet little is known about cheetahs’ resource use in areas where they co‐occur with people. The selection, or avoidance, of three anthropogenic variables (human footprint density, distance to main roads and wildlife areas) and five environmental variables (open habitat, semiclosed habitat, edge density, patch density and slope), at multiple spatial scales, was determined by analyzing collar data from six cheetahs. Cheetahs selected variables at different scales; anthropogenic variables were selected at broader scales (720–1440 m) than environmental variables (90–180 m), suggesting that anthropogenic pressures affect habitat selection at a home‐range level, whilst environmental variables influence site‐level habitat selection. Cheetah presence was best explained by human presence, wildlife areas, semiclosed habitat, edge density and slope. Cheetahs showed avoidance for humans and steep slopes and selected for wildlife areas and areas with high proportions of semiclosed habitat and edge density. Understanding a species’ resource requirements, and how these might be affected by humans, is crucial for conservation. Using a multiscale approach, we provide new insights into the habitat selection of a large carnivore living in a human‐wildlife landscape.     

Sharing the same slope: Behavioral responses of a threatened mesocarnivore to motorized and nonmotorized winter recreation

Winter recreation is a widely popular activity and is expected to increase due to changes in recreation technology and human population growth. Wildlife are frequently negatively impacted by winter recreation, however, through displacement from habitat, alteration of activity patterns, or changes in movement behavior. We studied impacts of dispersed and developed winter recreation on Canada lynx (Lynx canadensis) at their southwestern range periphery in Colorado, USA. We used GPS collars to track movements of 18 adult lynx over 4 years, coupled with GPS devices that logged 2,839 unique recreation tracks to provide a detailed spatial estimate of recreation intensity. We assessed changes in lynx spatial and temporal patterns in response to motorized and nonmotorized recreation, as well as differences in movement rate and path tortuosity. We found that lynx decreased their movement rate in areas with high‐intensity back‐country skiing and snowmobiling, and adjusted their temporal patterns so that they were more active at night in areas with high‐intensity recreation. We did not find consistent evidence of spatial avoidance of recreation: lynx exhibited some avoidance of areas with motorized recreation, but selected areas in close proximity to nonmotorized recreation trails. Lynx appeared to avoid high‐intensity developed ski resorts, however, especially when recreation was most intense. We conclude that lynx in our study areas did not exhibit strong negative responses to dispersed recreation, but instead altered their behavior and temporal patterns in a nuanced response to recreation, perhaps to decrease direct interactions with recreationists. However, based on observed avoidance of developed recreation, there may be a threshold of human disturbance above which lynx cannot coexist with winter recreation.     

基于野外痕迹点和GIS技术定量评估步道对大熊猫活动的影响

道路建设不仅直接导致野生动物死亡,还能对栖息地形成阻碍效应,导致小种群出现或隔离,增加物种灭绝的风险。生态学家在道路对野生动物影响研究中的一个重要进展是道路影响域(road-effectzone)的提出,但影响域既不能反映道路影响的变化性,也难以满足栖息地评估对数据的要求。为此,我们以大熊猫(Ailuropoda melanoleuca)为例来探讨道路影响的定量评估方法。在佛坪和长青保护区内选择了3条步道,获取了步道周边1,042个大熊猫的痕迹点数据,通过GIS计算各痕迹点到步道的距离,统计距离步道每20m内的痕迹点数量,以此作为其活动频率。在距步道每100m处设置检测点,通过非参数检验比较检测点前后活动频率分布的变化,寻找道路对大熊猫活动影响的突变点,确定影响变化的阈值距离和评价标准。研究发现在距离步道1,000m内,随距离的增加,大熊猫活动频率逐渐增大,大熊猫有明显的回避效应;在距步道500m、1,000m处发现活动频率发生了显著变化,为影响的阈值距离。本研究基于痕迹点和阈值距离的评估方法可以反映道路影响连续、渐变的特点,使定量、准确评估其影响成为可能。     

Hydrogeomorphology Influences Swamp Rabbit Habitat Selection in Bottomland Hardwood Forests

In the last century, bottomland hardwood (BLH) forests throughout the Lower Mississippi Alluvial Valley in the United States declined >80% and have been degraded because of habitat loss, fragmentation, and altered hydrology. To better understand how current conditions in BLH forest systems influence wildlife and to better manage land use and vegetation, we characterized winter (Dec–Mar) multi-scale habitat selection of 75 radio-marked swamp rabbits (Sylvilagus aquaticus) based on 850 locations in southern Illinois, USA, during 2010–2016. We investigated habitat selection by fitting resource selection functions with generalized linear mixed models based on Euclidean distances (km) to 8 cover types that described hydrogeomorphic conditions. At the second-order scale of selection (home range selection), swamp rabbits were closer to deciduous forest and low-elevation BLH and farther from agriculture, permanent water, shallow BLH, and woody wetland. At the third-order scale of selection (habitat selection within the home range), swamp rabbits selected areas closer to deciduous forest, low BLH, and shallow BLH, and farther from woody wetlands. For the swamp rabbit in Illinois, a BLH specialist at the northern extent of their range, habitat selection is limited to available terrestrial habitat that provides vegetation for food and hiding cover within linear and flood-prone BLH corridors surrounded by agricultural cover types that are largely unsuitable as habitat. Because hydrologic conditions are spatially and temporally dynamic, wildlife managers should focus on providing diverse habitat conditions across elevations that ensure the continuous availability of terrestrial habitat regardless of water level and flooding extent across the BLH landscape. Further reforestation efforts in BLH ecosystems should target current agricultural land on higher elevations adjacent to characteristically flood-prone forest remnants that escaped agricultural clearing due to frequent flooding. © 2021 The Wildlife Society.     

Return of Fire as a Restoration Tool: Long‐Term Effects of Burn Severity on Habitat Use by Mexican Fox Squirrels

After decades of suppression, fire is returning to forests of the western United States through wildfires and prescribed burns. These fires may aid restoration of vegetation structure and processes, which could improve conditions for wildlife species and reduce severe wildfire risk. Understanding response of wildlife species to fires is essential to forest restoration because contemporary fires may not have the same effects as historical fires. Recent fires in the Chiricahua Mountains of southeastern Arizona provided opportunity to investigate long‐term effects of burn severity on habitat selection of a native wildlife species. We surveyed burned forest for squirrel feeding sign and related vegetation characteristics to frequency of feeding sign occurrence. We used radio‐telemetry within fire‐influenced forest to determine home ranges of Mexican fox squirrels, Sciurus nayaritensis chiricahuae, and compared vegetation characteristics within home ranges to random areas available to squirrels throughout burned conifer forest. Squirrels fed in forest with open understory and closed canopy cover. Vegetation within home ranges was characterized by lower understory density, consistent with the effects of low‐severity fire, and larger trees than random locations. Our results suggest that return of low‐severity fire can help restore habitat for Mexican fox squirrels and other native wildlife species with similar habitat affiliations in forests with a historical regime of frequent, low‐severity fire. Our study contributes to an understanding of the role and impact of fire in forest ecosystems and the implications for forest restoration as fire returns to the region.     

Effects of Recreation and Hunting on Flushing Distance of Capercaillie

Abstract: The capercaillie (Tetrao urogallus) in Central Europe is an endangered species of grouse that is thought to be highly susceptible to human disturbance, possibly causing local populations to decline. We investigated the behavioral response of capercaillie in the Black Forest, Germany, and the French Pyrenees to an off-trail hiker by measuring flushing distances. Flushing distance varied with capercaillie sex, visibility of the hiker, intensity of winter tourism, and hunting pressure. Independent of the study area, males flushed at consistently longer distances than females, and lower visual blocking between bird and hiker resulted in longer flushing distances. Capercaillie flew at longer distances from an approaching hiker in areas with high intensity of winter tourism or hunting pressure than in undisturbed areas. We recommend the establishment of regulations requiring hikers to stay on trails and to close trails where intertrail distances fall below 100 m (90% of all flushing events appeared within 50 m). Furthermore, planting or preserving evergreen conifer trees in dense rows along critical parts of disturbance sources, reducing the degree of visibility between capercaillie and recreationists would increase habitat for capercaillie in forests with predictable recreation activities.     

Quantifying spatial habitat loss from hydrocarbon development through assessing habitat selection patterns of mule deer

Extraction of oil and natural gas (hydrocarbons) from shale is increasing rapidly in North America, with documented impacts to native species and ecosystems. With shale oil and gas resources on nearly every continent, this development is set to become a major driver of global land‐use change. It is increasingly critical to quantify spatial habitat loss driven by this development to implement effective mitigation strategies and develop habitat offsets. Habitat selection is a fundamental ecological process, influencing both individual fitness and population‐level distribution on the landscape. Examinations of habitat selection provide a natural means for understanding spatial impacts. We examined the impact of natural gas development on habitat selection patterns of mule deer on their winter range in Colorado. We fit resource selection functions in a Bayesian hierarchical framework, with habitat availability defined using a movement‐based modeling approach. Energy development drove considerable alterations to deer habitat selection patterns, with the most substantial impacts manifested as avoidance of well pads with active drilling to a distance of at least 800 m. Deer displayed more nuanced responses to other infrastructure, avoiding pads with active production and roads to a greater degree during the day than night. In aggregate, these responses equate to alteration of behavior by human development in over 50% of the critical winter range in our study area during the day and over 25% at night. Compared to other regions, the topographic and vegetative diversity in the study area appear to provide refugia that allow deer to behaviorally mediate some of the impacts of development. This study, and the methods we employed, provides a template for quantifying spatial take by industrial activities in natural areas and the results offer guidance for policy makers, mangers, and industry when attempting to mitigate habitat loss due to energy development.     

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.     

The Influence of Snowmobile Trails on Coyote Movements during Winter in High-Elevation Landscapes

Competition between sympatric carnivores has long been of interest to ecologists. Increased understanding of these interactions can be useful for conservation planning. Increased snowmobile traffic on public lands and in habitats used by Canada lynx (Lynx canadensis) remains controversial due to the concern of coyote (Canis latrans) use of snowmobile trails and potential competition with lynx. Determining the variables influencing coyote use of snowmobile trails has been a priority for managers attempting to conserve lynx and their critical habitat. During 2 winters in northwest Wyoming, we backtracked coyotes for 265 km to determine how varying snow characteristics influenced coyote movements; 278 km of random backtracking was conducted simultaneously for comparison. Despite deep snow (>1 m deep), radio-collared coyotes persisted at high elevations (>2,500 m) year-round. All coyotes used snowmobile trails for some portion of their travel. Coyotes used snowmobile trails for 35% of their travel distance (random: 13%) for a mean distance of 149 m (random: 59 m). Coyote use of snowmobile trails increased as snow depth and penetrability off trails increased. Essentially, snow characteristics were most influential on how much time coyotes spent on snowmobile trails. In the early months of winter, snow depth was low, yet the snow column remained dry and the coyotes traveled off trails. As winter progressed and snow depth increased and snow penetrability increased, coyotes spent more travel distance on snowmobile trails. As spring approached, the snow depth remained high but penetrability decreased, hence coyotes traveled less on snowmobile trails because the snow column off trail was more supportive. Additionally, coyotes traveled closer to snowmobile trails than randomly expected and selected shallower snow when traveling off trails. Coyotes also preferred using snowmobile trails to access ungulate kills. Snow compaction from winter recreation influenced coyote movements within an area containing lynx and designated lynx habitat.     

Habitat selection by Eurasian lynx (Lynx lynx) is primarily driven by avoidance of human activity during day and prey availability during night

The greatest threat to the protected Eurasian lynx (Lynx lynx) in Central Europe is human‐induced mortality. As the availability of lynx prey often peaks in human‐modified areas, lynx have to balance successful prey hunting with the risk of encounters with humans. We hypothesized that lynx minimize this risk by adjusting habitat choices to the phases of the day and over seasons. We predicted that (1) due to avoidance of human‐dominated areas during daytime, lynx range use is higher at nighttime, that (2) prey availability drives lynx habitat selection at night, whereas high cover, terrain inaccessibility, and distance to human infrastructure drive habitat selection during the day, and that (3) habitat selection also differs between seasons, with altitude being a dominant factor in winter. To test these hypotheses, we analyzed telemetry data (GPS, VHF) of 10 lynx in the Bohemian Forest Ecosystem (Germany, Czech Republic) between 2005 and 2013 using generalized additive mixed models and considering various predictor variables. Night ranges exceeded day ranges by more than 10%. At night, lynx selected open habitats, such as meadows, which are associated with high ungulate abundance. By contrast, during the day, lynx selected habitats offering dense understorey cover and rugged terrain away from human infrastructure. In summer, land‐cover type greatly shaped lynx habitats, whereas in winter, lynx selected lower altitudes. We concluded that open habitats need to be considered for more realistic habitat models and contribute to future management and conservation (habitat suitability, carrying capacity) of Eurasian lynx in Central Europe.     

人类休闲活动对森林鸟类群落无有害影响

我们对西班牙马德里森林深处(未受到人类干扰)、森林小路沿线(间或有徒步旅行者经过)以及森林野餐区(有较持久干扰)的鸟类进行了调查,以查明人类休闲活动对该地区鸟类群落的分布和多度的影响.结合样方法对以上3类区域鸟类群落的总体特征(如:鸟类的多度、物种的丰富度和多样性)、不同鸟类群体的特征(如:鸦科鸟类的多度,营巢或者取食集团的特征)、各鸟种的多度等参数进行了比较.结果表明:小路沿线和野餐区样方内鸟类的多度和物种丰富度比森林深处更高,已查明14种鸟类的多度在人为干扰区域比森林深处高,只有5种鸟类的多度在森林深处更高一些;在树冠层筑巢的鸟类、在树干和地面觅食的鸟类及鸦科鸟类在休闲活动区有更高的多度;地巢性鸟类在休闲区的多度较低,这种差别尤以阔叶落叶林中加明显.而森林小路的鸟类群落结构与森林内部非常相似,说明人类休闲活动对这些林区鸟类群落的影响更多缘自野餐.原因是野餐区常常具有以下特点:具有高大的乔木、中层植被稀疏、有较多的洞和缝隙以及野餐后会遗留下一些垃圾.以上结论与研究的预期目标正好相反,人类的这些休闲活动可能会降低地巢性鸟类的适宜度,但是对大多数鸟的类栖息地没有不利影响.     

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.     

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.     

Habitat selection by an Alpine ungulate: the significance of forage characteristics varies with scale and season

Habitat selection in ungulates should ensure access to abundant forage of sufficiently high quality. Species living in rugged mountain areas have to face nutritional bottlenecks regularly and should show particularly sophisticated habitat selection behaviour. However, patterns and mechanisms of such adaptations remain little studied. We analysed habitat selection and its seasonal variability of 10 GPS‐collared red deer Cervus elaphus living in a topographically challenging landscape of the Swiss Alps. We hypothesised that resource selection by red deer was scale‐dependent and predicted that scale‐dependence would vary among seasons in relation to seasonal changes of available forage biomass and quality, which we sampled across the entire study area of 250 km². The studied population of Alpine red deer undertook altitudinal migrations and showed scale‐dependent habitat selection that was strongest in winter and declined through spring and summer. Selection occurred mostly at the larger (landscape/home‐range location) scale and less so at the smaller (within home‐range) scale. Topographic parameters were selected mainly at the landscape scale and mostly in winter. About 70% of all instances of preference for habitat parameters were associated with above‐average forage characteristics, represented mostly by higher crude protein content, in a few cases also by higher biomass or both. The overall pattern of space use by red deer characterised by migration and seasonal habitat selection was therefore closely linked to the quality of food resources, although some trade‐offs with avoiding human disturbance may also have been involved.     

Red deer (<Emphasis Type="Italic">Cervus elaphus</Emphasis>) avoid mountain biking trails

Human outdoor activities commonly affect animal behaviour. Ungulates often avoid roads and trails and increase their avoidance with expanding and intensifying human recreational activity. Recently, mountain biking has become increasingly popular in many regions in Norway, but we still have limited knowledge about how mountain biking may affect wildlife. In this study, we used pellet-group counts and camera traps to study the effect of mountain biking on red deer abundance in Kaupanger, Norway. Pellet-group counts decreased close to biking trails, and it appeared that red deer avoided using areas within 40 m of such trails. Camera trap data showed that there was a tendency for deer abundance to decrease with increasing human activity (trail width) during daytime. Males reacted more strongly to this increasing activity than did females. Even with a small-scale approach, focusing on within-stand effects, we found that red deer were affected by mountain bikers and avoided areas close to biking trails. Our findings imply that any further increase in mountain biking may result in greater avoidance and, thus, less suitable habitat for red deer in forest areas.     

Nonmotorized recreation and motorized recreation in shrub‐steppe habitats affects behavior and reproduction of golden eagles (Aquila chrysaetos)

Different forms of outdoor recreation have different spatiotemporal activity patterns that may have interactive or cumulative effects on wildlife through human disturbance, physical habitat change, or both. In western North America, shrub‐steppe habitats near urban areas are popular sites for motorized recreation and nonmotorized recreation and can provide important habitat for protected species, including golden eagles. Our objective was to determine whether recreation use (i.e., number of recreationists) or recreation features (e.g., trails or campsites) predicted golden eagle territory occupancy, egg‐laying, or the probability a breeding attempt resulted in ≥1 offspring (nest survival). We monitored egg‐laying, hatching and fledging success, eagle behavior, and recreation activity within 23 eagle territories near Boise, Idaho, USA. Territories with more off‐road vehicle (ORV) use were less likely to be occupied than territories with less ORV use (β = ?1.6, 85% CI: ?2.8 to ?0.8). At occupied territories, early season pedestrian use (β = ?1.6, 85% CI: ?3.8 to ?0.2) and other nonmotorized use (β = ?3.6, 85% CI: ?10.7 to ?0.3) reduced the probability of egg‐laying. At territories where eagles laid eggs, short, interval‐specific peaks in ORV use were associated with decreased nest survival (β = ?0.5, 85% CI: ?0.8 to ?0.2). Pedestrians, who often arrived near eagle nests via motorized vehicles, were associated with reduced nest attendance (β = ?11.9, 85% CI: ?19.2 to ?4.5), an important predictor of nest survival. Multiple forms of recreation may have cumulative effects on local populations by reducing occupancy at otherwise suitable territories, decreasing breeding attempts, and causing nesting failure. Seasonal no‐stopping zones for motorized vehicles may be an alternative to trail closures for managing disturbance. This study demonstrates the importance of considering human disturbance across different parts of the annual cycle, particularly where multiple forms of recreation have varying spatiotemporal use patterns that create human–wildlife interactions.     

Roe deer at risk: teasing apart habitat selection and landscape constraints in risk exposure at multiple scales

Non‐consumptive effects of predators result from the cost of responses to perceived risk. Prey modulate risk exposure through flexible habitat selection at multiple scales which, in interaction with landscape constraints, determines their use of risky habitats. Identifying the relative contributions of landscape constraints and habitat selection to risk exposure is a critical first step towards a mechanistic understanding of non‐consumptive effects. Here, we provide an integrative multi‐scale study of roe deer spatial responses to variable hunting pressure along a landscape gradient of open habitats and dispersed refuges. Between low‐risk and high‐risk periods, we investigated shifts in 1) home‐range location, 2) probability of using risky habitats (between‐habitat scale) and 3) distance to the nearest refuge (within‐habitat scale). For 2) and 3), we disentangled the contributions of landscape constraints and habitat selection to risky habitat use. We found that when risk was high, roe deer did not shift their home‐range, but generally decreased their use of risky habitats, and sometimes reduced their distance to cover (particularly older animals). There was a functional response in between‐habitat selection, with animals living in more open landscapes responding more than those living in landscapes with more refuges. However, individuals living in more open landscapes avoided open risky habitat less. Finally, we found that among‐individual variation in risk exposure was generally, but not always, minimized by habitat selection across gradients of landscape constraints. To our knowledge, this is the first study simultaneously documenting prey responses to risk at the within‐habitat, between‐habitat and home‐range scales. Our results support the view that between‐habitat selection acts at a higher hierarchical level than within‐habitat selection, and provide a framework for disentangling the contributions of habitat selection and landscape constraints to risk exposure. Selection cannot always compensate for landscape constraints, indicating a need for further investigation of the processes underlying habitat selection.     

Effects of winter recreation on northern ungulates with focus on moose (Alces alces) and snowmobiles

Winter recreation can displace ungulates to poor habitats, which may raise their energy expenditure and lower individual survivorship, causing population declines. Winter recreation could be benign, however, if animals habituate. Moreover, recreation creates trails. Traveling on them could reduce energy expenditure, thereby increasing ungulate survivorship and generating population benefits. Balancing recreation use with wildlife stewardship requires identifying when these effects occur. This task would be simpler if guidelines existed to inform assessments. We developed and tested such guidelines using two approaches. First, we synthesized literature describing the effects of winter recreation—motorized and nonmotorized—on northern ungulates. This synthesis enabled formulating six guidelines, while exposing two requiring further attention (ungulate habituation and displacement). Second, we tested these two guidelines and evaluated the others by quantifying the behavioral responses of moose to snowmobiles, in two areas of south-central Alaska, differing by snowmobile predictability. For each location, we modeled moose preferences during the snowmobile period using different combinations of eight variables—static (elevation and slope), biotic (habitat and cover), and anthropogenic (distance to roads, railroads, snowmobile trails, and trail density). We identified the model with the most support and used it to estimate parameter coefficients for pre- and post-recreation periods. Changes in coefficients between periods indicated snowmobile effects on moose. Overall, we produced and evaluated six guidelines describing when winter recreation is potentially detrimental to ungulates as follows: (1) when unpredictable, (2) spanning large areas, (3) long in duration, (4) large spatial footprint, (5) nonmotorized, and (6) when animals are displaced to poor quality habitats.     

Seasonal habitat selection and space use by a semi‐free range herbivore in a heterogeneous savanna landscape

Understanding factors that influence habitat selection in heterogeneous landscapes is fundamental for establishing realistic models on animal distribution to inform rangeland management. In this study, we tested whether seasonal variation in habitat selection within the home range of a large herbivore was influenced by constraints such as, distances from water and central place using semi‐free range cattle (Bos taurus) as a case study. We also tested whether shifts in space use over time were dependent on spatial scale and on the overall abundance of resources. We predicted that distance from water significantly influenced dry season habitat selection while the influence of the central place on habitat selection was season‐independent. We also predicted that shifts in space use over time were spatial scale‐dependent, and that large herbivores would include more diverse habitats in their home ranges during the dry season, when water and food resources are less abundant. Multinomial logit models were used to construct habitat selection models with distances from water and central place as habitat‐specific constraints. Results showed significant variations in habitat selection between the dry and wet season. As predicted, the effect of distance from central place was season‐independent, while the effect of water was not included in the top dry season models contrary to expectation. A diverse range of habitats were also selected during the dry season including agricultural fields. Results also indicated that shifts in space use were spatial scale dependent, with core areas being more sensitive to changes than the home range. In addition, shifts in space use responded to temporal changes in habitat composition. Overall, our results suggest that semi‐free range herbivores adopt different foraging strategies in response to spatial‐temporal changes in habitat availability.     

Ecological correlates of Himalayan musk deer Moschus leucogaster

Himalayan musk deer (Moschus leucogaster; hereafter musk deer) are endangered as a result of poaching and habitat loss. The species is nocturnal, crepuscular, and elusive, making direct observation of habitat use and behavior difficult. However, musk deer establish and repeatedly use the same latrines for defecation. To quantify musk deer habitat correlates, we used observational spatial data based on presence–absence of musk deer latrines, as well as a range of fine spatial‐scale ecological covariates. To determine presence–absence of musk deer, we exhaustively searched randomly selected forest trails using a 20‐m belt transect in different study sites within the Neshyang Valley in the Annapurna Conservation Area. In a subsequent way, study sites were classified as habitat or nonhabitat for musk deer. A total of 252 plots, 20 × 20 m, were systematically established every 100 m along 51 transects (each ~0.5 km long) laid out at different elevations to record a range of ecological habitat variables. We used mixed‐effect models and principal component analysis to characterize relationships between deer presence–absence data and habitat variables. We confirmed musk deer use latrines in forests located at higher elevations (3,200–4,200 m) throughout multiple seasons and years. Himalayan birch (Betula utilis) dominated forest, mixed Himalayan fir (Abies spectabilis), and birch forest were preferred over pure Himalayan fir and blue pine (Pinus wallichiana) forest. Greater crown cover and shrub diversity were associated with the presence of musk deer whereas tree height, diameter, and diversity were weakly correlated. Topographical attributes including aspect, elevation, distance to water source, and slope were also discriminated by musk deer. Over‐ and understory forest management can be used to protect forests likely to have musk deer as predicted by the models to ensure long‐term conservation of this rare deer.