Influence of Well Pad Activity on Winter Habitat Selection Patterns of Mule Deer

ABSTRACT Conversion of native winter range into producing gas fields can affect the habitat selection and distribution patterns of mule deer (Odocoileus hemionus). Understanding how levels of human activity influence mule deer is necessary to evaluate mitigation measures and reduce indirect habitat loss to mule deer on winter ranges with natural gas development. We examined how 3 types of well pads with varying levels of vehicle traffic influenced mule deer habitat selection in western Wyoming during the winters of 2005–2006 and 2006–2007. Well pad types included producing wells without a liquids gathering system (LGS), producing wells with a LGS, and well pads with active directional drilling. We used 36,699 Global Positioning System locations collected from a sample (n = 31) of adult (>1.5-yr-old) female mule deer to model probability of use as a function of traffic level and other habitat covariates. We treated each deer as the experimental unit and developed a population-level resource selection function for each winter by averaging coefficients among models for individual deer. Model coefficients and predictive maps for both winters suggested that mule deer avoided all types of well pads and selected areas further from well pads with high levels of traffic. Accordingly, impacts to mule deer could probably be reduced through technology and planning that minimizes the number of well pads and amount of human activity associated with them. Our results suggested that indirect habitat loss may be reduced by approximately 38–63% when condensate and produced water are collected in LGS pipelines rather than stored at well pads and removed via tanker trucks. The LGS seemed to reduce long-term (i.e., production phase) indirect habitat loss to wintering mule deer, whereas drilling in crucial winter range created a short-term (i.e., drilling phase) increase in deer disturbance and indirect habitat loss. Recognizing how mule deer respond to different types of well pads and traffic regimes may improve the ability of agencies and industry to estimate cumulative effects and quantify indirect habitat losses associated with different development scenarios.     

Behavioral and Demographic Responses of Mule Deer to Energy Development on Winter Range

Anthropogenic habitat modification is a major driver of global biodiversity loss. In North America, one of the primary sources of habitat modification over the last 2 decades has been exploration for and production of oil and natural gas (hydrocarbon development), which has led to demographic and behavioral impacts to numerous wildlife species. Developing effective measures to mitigate these impacts has become a critical task for wildlife managers and conservation practitioners. However, this task has been hindered by the difficulties involved in identifying and isolating factors driving population responses. Current research on responses of wildlife to development predominantly quantifies behavior, but it is not always clear how these responses scale to demography and population dynamics. Concomitant assessments of behavior and population-level processes are needed to gain the mechanistic understanding required to develop effective mitigation approaches. We simultaneously assessed the demographic and behavioral responses of a mule deer population to natural gas development on winter range in the Piceance Basin of Colorado, USA, from 2008 to 2015. Notably, this was the period when development declined from high levels of active drilling to only production phase activity (i.e., no drilling). We focused our data collection on 2 contiguous mule deer winter range study areas that experienced starkly different levels of hydrocarbon development within the Piceance Basin. We assessed mule deer behavioral responses to a range of development features with varying levels of associated human activity by examining habitat selection patterns of nearly 400 individual adult female mule deer. Concurrently, we assessed the demographic and physiological effects of natural gas development by comparing annual adult female and overwinter fawn (6-month-old animals) survival, December fawn mass, adult female late and early winter body fat, age, pregnancy rates, fetal counts, and lactation rates in December between the 2 study areas. Strong differences in habitat selection between the 2 study areas were apparent. Deer in the less-developed study area avoided development during the day and night, and selected habitat presumed to be used for foraging. Deer in the heavily developed study area selected habitat presumed to be used for thermal and security cover to a greater degree. Deer faced with higher densities of development avoided areas with more well pads during the day and responded neutrally or selected for these areas at night. Deer in both study areas showed a strong reduction in use of areas around well pads that were being drilled, which is the phase of energy development associated with the greatest amount of human presence, vehicle traffic, noise, and artificial light. Despite divergent habitat selection patterns, we found no effects of development on individual condition or reproduction and found no differences in any of the physiological or vital rate parameters measured at the population level. However, deer density and annual increases in density were higher in the low-development area. Thus, the recorded behavioral alterations did not appear to be associated with demographic or physiological costs measured at the individual level, possibly because populations are below winter range carrying capacity. Differences in population density between the 2 areas may be a result of a population decline prior to our study (when development was initiated) or area-specific differences in habitat quality, juvenile dispersal, or neonatal or juvenile survival; however, we lack the required data to contrast evidence for these mechanisms. Given our results, it appears that deer can adjust to relatively high densities of well pads in the production phase (the period with markedly lower human activity on the landscape), provided there is sufficient vegetative and topographic cover afforded to them and populations are below carrying capacity. The strong reaction to wells in the drilling phase of development suggests mitigation efforts should focus on this activity and stage of development. Many of the wells in this area were directionally drilled from multiple-well pads, leading to a reduced footprint of disturbance, but were still related to strong behavioral responses. Our results also indicate the likely value of mitigation efforts focusing on reducing human activity (i.e., vehicle traffic, light, and noise). In combination, these findings indicate that attention should be paid to the spatial configuration of the final development footprint to ensure adequate cover. In our study system, minimizing the road network through landscape-level development planning would be valuable (i.e., exploring a maximum road density criteria). Lastly, our study highlights the importance of concomitant assessments of behavior and demography to provide a comprehensive understanding of how wildlife respond to habitat modification. © 2021 The Wildlife Society.     

Mule deer and energy development—Long‐term trends of habituation and abundance

As the extent and intensity of energy development in North America increases, so do disturbances to wildlife and the habitats they rely upon. Impacts to mule deer are of particular concern because some of the largest gas fields in the USA overlap critical winter ranges. Short‐term studies of 2–3 years have shown that mule deer and other ungulates avoid energy infrastructure; however, there remains a common perception that ungulates habituate to energy development, and thus, the potential for a demographic effect is low. We used telemetry data from 187 individual deer across a 17‐year period, including 2 years predevelopment and 15 years during development, to determine whether mule deer habituated to natural gas development and if their response to disturbance varied with winter severity. Concurrently, we measured abundance of mule deer to indirectly link behavior with demography. Mule deer consistently avoided energy infrastructure through the 15‐year period of development and used habitats that were an average of 913 m further from well pads compared with predevelopment patterns of habitat use. Even during the last 3 years of study, when most wells were in production and reclamation efforts underway, mule deer remained >1 km away from well pads. The magnitude of avoidance behavior, however, was mediated by winter severity, where aversion to well pads decreased as winter severity increased. Mule deer abundance declined by 36% during the development period, despite aggressive onsite mitigation efforts (e.g. directional drilling and liquid gathering systems) and a 45% reduction in deer harvest. Our results indicate behavioral effects of energy development on mule deer are long term and may affect population abundance by displacing animals and thereby functionally reducing the amount of available habitat.     

Temporal variation in habitat selection breaks the catch‐22 of spatially contrasting predation risk from multiple predators

Predator avoidance depends on prey being able to discern how risk varies in space and time, but this is made considerably more complicated if risk is simultaneously present from multiple predators. This is the situation for an increasing number of mammalian prey species, as large carnivores recover or are reintroduced in ecosystems on several continents. Roe deer Capreolus capreolus in southern Norway illustrate a case in which prey face two predators with contrasting patterns of predation risk. They face a catch‐22 situation: spatially avoiding the risk from one predator (lynx Lynx lynx in dense habitat) implies exposure to the other (hunters in open habitat). Using GPS‐data from 29 roe deer, we tested for daily and seasonal variation in roe deer selection for habitat with respect to the habitats’ year‐round average risk level. Generally, roe deer altered their habitat selection between night and day in a pattern consistent with being able to avoid predicted risk from the nocturnal lynx during night and predicted risk from human hunters during day. However, seasonal variation in habitat selection only partially corresponded with the predicted seasonal variation in risk. Whereas roe deer avoided areas with high risk from hunters more strongly during the hunting season than in other seasons, there was a lack of selection towards areas and time periods lowering the risk of lynx predation during winter. It seems likely that the risk of starvation and thermal stress constrain roe deer habitat selection during this energetically challenging season with cold temperatures, snow cover and limited natural forage. The habitat selection pattern of roe deer fits thus only partly with the two contrasting risk gradients they face. Adjusting risk‐avoidance behavior temporally can be an adaptive response in the case of several predators whose predation patterns differ in space and time.     

Migratory Disturbance Thresholds with Mule Deer and Energy Development

Fine-scale movement data has transformed our knowledge of ungulate migration ecology and now provides accurate, spatially explicit maps of migratory routes that can inform planning and management at local, state, and federal levels. Among the most challenging land use planning issues has been developing energy resources on public lands that overlap with important ungulate habitat, including the migratory routes of mule deer (Odocoileus hemionus). We generally know that less development is better for minimizing negative effects and maintaining habitat function, but we lack information on the amount of disturbance that animals can tolerate before reducing use of or abandoning migratory habitat. We used global positioning system data from 56 deer across 15 years to evaluate how surface disturbance from natural gas well pads and access roads in western Wyoming, USA, affected habitat selection of mule deer during migration and whether any disturbance threshold(s) existed beyond which use of migratory habitat declined. We used resource and step selection functions to examine disturbance thresholds at 3 different spatial scales. Overall, migratory use by mule deer declined as surface disturbance increased. Based on the weight of evidence from our 3 independent but complementary metrics, declines in migratory use related to surface disturbance were non-linear, where migratory use sharply declined when surface disturbance from energy development exceeded 3%. Disturbance thresholds may vary across regions, species, or migratory habitats (e.g., stopover sites). Such information can help with management and land use decisions related to mineral leasing and energy development that overlap with the migratory routes of ungulates. © 2020 The Wildlife Society.     

Spatial and temporal habitat use and selection by red deer: The use of direct and indirect methods

Given the importance of red deer Cervus elaphus for hunting and conservation purposes, understanding the interactions between this species and its habitats in the Mediterranean region is a crucial step for the sustainable management of this species. Aiming to compare pellet group counts and direct observations methods to study the habitat use and selection by red deer, the results obtained by both methods were compared, and their advantages and disadvantages were discussed. To understand the temporal patterns of habitat use and selection, the survey was conducted at three different seasons, birth period, rut season and winter. The habitat use and selection were studied in relation to land cover, watercourse, roads, ecotone zones and other topographic features (altitude, slope and aspect), using generalized linear models and selection ratios. The similarity of the results provided by pellet group counts and direct observations indicate that both methods may constitute useful tools to study the habitat use and selection by red deer. Globally, red deer seemed to select habitats that provide simultaneously food and some cover, as shown by its preference for shrublands, independently of the sampling season. The positive selection of ecotone zones embodies the need for open spaces. Males and females showed a similar use of shrubland, but selected patches with different characteristics therein. The spatial and temporal patterns exhibited by our results suggest that red deer balance their habitat requirements in respect to each phase of their reproductive cycle. Pellet group counts and direct observations seem to be useful methods to analyze habitat use and selection, and may provide helpful knowledge to the management and conservation of red deer.     

Long-term density-dependent changes in habitat selection in red deer (Cervus elaphus)

Understanding how habitat selection changes with population density is a key concept in population regulation, community composition and managing impacts on biodiversity and ecosystem services. At low density, it is expected that individuals select habitats in terms of their preference, but as population density increases, the availability of resources per individual declines on preferred habitats, leading to competition which forces some individuals to exploit less preferred habitats. Using spatial information of Scottish red deer (Cervus elaphus) winter counts, carried out in 110 areas across Scotland between 1961 and 2004 (a total of 1,206,495 deer observations), we showed how winter habitat niche breadth in red deer has widened with increasing population density. Heather moorland and montane habitats were most and least preferred for deer, respectively. Increasing density favoured the selection of grassland, to the detriment of the selection of heather moorland. The selection of heather and grassland decreased when temperature increased, while the selection of montane and peatland habitats increased. These findings are important for understanding how habitat use, density and population are likely to be affected by weather, and allow us to predict habitat impacts by large mammal herbivory and climate.     

So close and yet so different: The importance of considering temporal dynamics to understand habitat selection

Habitat selection of animals is influenced by spatial heterogeneity as well as temporal environmental dynamics. In addition, human activities potentially have severe influences on the habitat selection of animals, often resulting in more nocturnal behavior. We investigated seasonal and circadian habitat selection patterns of red deer (Cervus elaphus) on a military training area in Bavaria (Germany). Individual deer ranged on two neighboring, but non-overlapping sites differing in landscape composition and human activity. Using GPS telemetry data, we visually investigated selection patterns and then fitted step-selection functions to assess multiple approaches to account for temporal (i.e., diel and seasonal) effects and for the impact of habitat conditions on selection. We first showed that the way in which time of day is considered in step-selection functions is essential for obtaining ecologically meaningful results, and that the usual ways of including time (i.e., either based on clock-time or based on night vs. day categories) can lead to incomplete or misleading conclusions. Furthermore, we found that individuals followed either circadian or seasonal habitat selection patterns, depending on the site they inhabited. This can be explained by differences in disturbance between the two sites. Except in winter, individuals selected for open habitats during the night irrespective of human activity level, but only individuals from the more disturbed site selected for covered habitats during the day. Our results highlight the importance of carefully considering the appropriate temporal scale for habitat selection analyses. Our findings also indicate that red deer are not a crepuscular or nocturnal species per se, as is often observed in human-dominated landscapes. Instead, our results imply that nocturnality in red deer is an effect of adaption to human activities.     

Red deer habitat selection and movements in relation to roads

Roads affect wildlife in many direct and indirect ways. For ungulates, roads may inhibit seasonal migration and may cause an effective loss of habitat due to avoidance. On the other hand, roadsides and associated agricultural lands offer high quality forage that may attract ungulates and increase the frequency of car accidents. Mitigating actions require detailed knowledge on space use in relation to roads. Using data from 67 global positioning system (GPS)-marked red deer in Norway, we quantified 1) scale of avoidance of roads, 2) crossing frequency, and 3) selection of crossing sites. Red deer avoided roads only on a very local scale and only during daytime, with minor influence of variation in road size (traffic burden). Marked red deer crossed roads, on average, 2 times per day. Females crossed more frequently than males and crossings were most frequent during autumn and winter and during night. Deer selected forested crossing sites close to agricultural pastures, reflecting that roads are crossed most often on nightly feeding excursions. Our findings imply that red deer in our study area have adjusted to exploit feeding habitat close to roads at times of low traffic burden. The high frequency of crossings suggests a limited influence on seasonal migration patterns. The frequency at which red deer cross highways suggests that mitigation measures to reduce road mortality may be effective if targeted in the right areas. © 2012 The Wildlife Society.     

Experimental evidence that deer browsing reduces habitat suitability for breeding Common Nightingales Luscinia megarhynchos

The ecological impacts of increasing populations of deer (Cervidae) in Europe and North America are becoming more widespread and pronounced. Within Britain, it has been suggested that declines in several woodland bird species, particularly those dependent on dense understorey vegetation, may be at least partly due to these effects. Here we present experimental evidence of the effects of deer browsing on the fine‐scale habitat selection and habitat use by a bird species in Europe. The study was conducted in a wood in eastern England where a decrease in Common Nightingale Luscinia megarhynchos numbers has coincided with a large increase in deer numbers. Eight woodland plots were cut to produce young coppice regrowth (a favoured habitat for Nightingales). Deer were excluded from half of each plot using steel fences, thus creating eight experimental pairs of exclosures (unbrowsed) and controls (browsed). Radiotelemetry and territory mapping of male Nightingales showed strong selection of exclosures. The density of territories was 15 times greater in the exclosures than in grazed controls. Selection for exclosures was significant for the minimum convex polygon, 95% kernel and 50% core home‐ranges used by seven radiotracked males. Tracked birds spent 69% of their time in the 6% of the study area protected from deer. Intensified browsing by deer influenced local settlement patterns of Nightingales, supporting the conclusion that increased deer populations are likely to have contributed to declines of Nightingales in Britain, and potentially those of other bird species dependent on dense understorey.     

Migratory strategies, fawn recruitment, and winter habitat use by urban and rural mule deer (Odocoileus hemionus)

We examined the migratory strategies of mule deer (Odocoileus hemionus) using adjacent urban and rural winter ranges in northern Utah in relation to deer demography and patterns of habitat use. Urban deer were more likely to be migratory than rural deer, even though migratory animals from the two herds intermixed on a common, high-elevation summer range. Urban deer exhibited lower fawn recruitment than rural deer; but within each herd, demographic characteristics of migratory and nonmigratory animals suggested that game theory explained the ratios of deer adopting each behavior. Estimates of animal numbers and available habitat did not reveal clearly whether deer densities differed on the two winter ranges. However, patterns of habitat use by urban deer were so clustered around areas of concealment vegetation that animals probably experienced higher local densities than rural animals. In addition, the clustered patterns of habitat use by urban deer resulted in incomplete use of available forage. This may have contributed to the relatively poor fawn recruitment by urban deer, a phenomenon that appeared to be perpetuated by their strong fidelity to winter range.     

Thermal constraints and optimization of winter feeding and habitat choice in white-tailed deer

I examined the activity and habitat choice of white-tailed deer in winter to determine how deer balance feeding in habitats which may expose them to cold with resting in sheltered habitats. Average heat gain or loss of deer for feeding, resting and walking in four habitats in each of 24 h were estimated for two winter periods (early-mid and late winter). These values were used in a dynamic optimization model to predict the hourly behavior-habitat choices which enable deer to maintain thermal homeostasis, minimize the risk of exceeding lethal body temperatures and satisfy daily energy requirements. The behavior and habitat choices observed under natural conditions were consistent with the model predictions. Deer foraged for the maximum time that was thermally possible in both sampling periods and they used exposed habitats without incurring high heat losses. Deer appeared to avoid overheating in sheltered habitats during daylight and avoid hypothermia in exposed habitats at night.     

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.     

Linking alternative food sources to winter habitat selection of herbivores in overbrowsed landscapes

During winter, ungulates in boreal forests must cope with high energetic costs related to locomotion in deep snow and reduced forage abundance and quality. At high density, ungulates face additional constraints, because heavy browsing reduces availability of woody browse, the main source of forage during winter. Under these severe conditions, large herbivores might forage on alternative food sources likely independent of browsing pressure, such as litterfall or windblown trees. We investigated the influence of alternative food sources on winter habitat selection, by studying female white-tailed deer (Odocoileus virginianus) living in 2 landscapes with contrasted browse abundance, recently logged and regenerated landscapes, in a population at high density and on a large island free of predators. We fitted 21 female white-tailed deer with Global Positioning System (GPS) collars and delineated winter home ranges and core areas. We measured snow conditions in different habitat categories and sampled vegetation in the core areas and in the rest of the home ranges to determine how forage abundance, protective cover, and snow conditions influenced habitat selection within the home range. In both landscapes, deer were less likely to use open habitat categories as snow accumulated on the ground. At a finer scale, deer inhabiting the regenerated landscape intensively used areas where balsam fir cover was intermediate with greater balsam fir browse density than in the rest of the home range. In the recently logged landscape, deer were more likely to be found near edges between clear-cuts and balsam fir stands and in areas where windblown balsam fir trees were present; the latter being the most influential variable. Although balsam fir browse was sparse and mainly out of reach in this landscape, deer increased the use of areas where it was present. Our results offer novel insights into the resource selection processes of northern ungulates, as we showed that access to winter forage, such as woody browse and alternative food sources, depends on climatic conditions and stochastic events, such as abundant compacted snow or windthrows. To compensate for these scarce and unpredictable food supplies, deer selected habitat categories, but mostly areas within those habitat categories, where the likelihood of finding browse, litterfall, and windblown trees was greatest. © 2011 The Wildlife Society.     

察青松多白唇鹿国家级自然保护区白唇鹿对夏季生境的选择

白唇鹿是青藏高原特有鹿种,人们对其生境选择与利用知之甚少。2009 年6 -8 月和2011 年7 - 8 月,用痕迹检验法和直接观察法研究了察青松多白唇鹿国家级自然保护区内白唇鹿的夏季生境选择。在保护区内若当沟和麻绒沟两个白唇鹿集中分布区域测定了195 个样方中的植被类型、海拔高度、坡位、坡度、坡向、离水源距离、隐蔽度和人类活动干扰8 个生态因子。用Vanderloeg 和Scavia 选择指数和主成分分析法分析了白唇鹿夏季生境选择的影响因子。结果表明:白唇鹿夏季倾向于选择海拔4 000 m 以上中上坡位的高山草甸生境和离水源距离500 m 以外的阳坡或阴坡生境;而回避坡度大于60 度、裸岩生境、下坡位生境和半阴坡生境。除隐蔽条件和人类活动干扰外,其余6 个生态因子均对白唇鹿夏季生境选择有明显的影响。生态因子主成分分析表明前3 个主成分的贡献量为68. 8% ,其中:第1 主成分的贡献率为39.76% ,该主成分与坡位、坡度和海拔的相关系数绝对值较大;第二主成分的贡献率为16.32% ,与其相关系数绝对值较高的是隐蔽条件与植被类型;第三主成分的贡献率为12. 71% ,与其相关系数绝对值较高的是坡向。     

Temporal plasticity in thermal‐habitat selection of burbot Lota lota a diel‐migrating winter‐specialist

In this study, animal‐borne telemetry with temperature sensors was coupled with extensive habitat temperature monitoring in a dimictic reservoir, to test the following hypotheses: behavioural thermoregulation occurs throughout the year and temperature selection varies on a diel and seasonal basis, in a winter‐specialist diel‐migrating fish. Burbot Lota lota demonstrated nightly behavioural thermoregulation throughout the year, with a large seasonal shift between selection for very cold temperatures (<2° C) optimal for reproduction during the spawning period and selection for warmer temperatures (12–14° C) optimal for hunting and feeding during non‐reproductive periods. During daylight hours, while L. lota avoided habitats warmer than optimal for reproduction and feeding during the spawning and non‐reproductive periods, respectively, active selection was limited to selection for 4–6° C habitat during the prespawning period. Although behavioural thermoregulation explained the night‐time migration, behavioural thermoregulation only partially explained daytime behaviour, indicating that diel migration is best explained by a combination of factors. Thus, thermal‐habitat selection was a good predictor of night‐time habitat occupancy in a diel‐migrating species. Together, these results show that thermal‐habitat selection by fishes may be important throughout the year and a more seasonally plastic behaviour than previously recognized.     

老爷岭南部狍冬季移动、卧息生境选择及其适宜性评价

在动物生境研究中,移动生境和卧息生境是生境研究的焦点。开展移动生境和卧息生境选择,并在此基础上进行生境评价,有利于深入了解动物对移动和卧息生境条件的需求,制定科学合理的栖息地保护计划。以东北虎（Panthera tigris altaica）的主要猎物物种之一-狍（Capreolus pygargus）为研究对象,于2017-2019年冬季积雪覆盖期在老爷岭南部通过随机布设28个大样方和84条用于足迹链跟踪的样线收集狍的移动点和卧息点信息,再结合近年来收集的东北虎出现点,利用广义可加模型（GAM）和最大熵模型（MaxEnt）进行狍移动、卧息生境选择及评价研究。移动生境选择研究表明,狍在移动的过程中偏好选择坡度小、距农田距离>500 m、远离道路、居民点和低海拔或较高海拔的区域;移动生境评价分析表明,移动适宜和次适宜生境面积之和为1318.16 km²,占研究区域面积的51.28%,当加入虎活动点影响因子后,狍移动适宜和次适宜生境面积之和为901.52 km²,适宜和次适宜生境面积之和减少了31.61%。狍卧息生境选择研究表明,水源、农田、道路和雪深是影响狍卧息的关键因素,其中雪深对狍卧息生境选择的贡献率达到70.13%;卧息生境评价表明,卧息适宜和次适宜生境面积之和为1243.77 km²,占研究区域面积的48.39%,当加入虎出现点因子后,适宜生境和次适宜生境面积之和减少了61.00%,仅为485.02 km²。研究认为,虎的出现对狍移动和卧息生境选择均产生影响,虎的活动及捕食行为可能会减少狍的活动范围和频次,狍远离虎活动区域卧息休息,压缩了狍适宜卧息的空间。     

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.     

Experimental evidence that competition and habitat use shape the individual fitness surface

A key prediction made by theories of density‐dependent competition is that resource overlap should increase the intensity of competition. By extension, we can predict that competition should lead to density‐dependent natural selection. I studied natural selection on limb length and body size in a total of seven populations of Anolis sagrei over 3 years in the Bahamas. Experimental manipulations of population density on small off‐shore cays revealed that the strength of natural selection on body size increased with density, suggesting that density‐dependent intraspecific competition drives natural selection. At low density, reduced competition revealed significant selection on limb length driven by changes in perch diameter, indicating that selection favoured a match between morphology and habitat. The role habitat played in shaping selection was further illuminated by inter‐annual changes in vegetation structure stemming from variation in precipitation among years. Thus, changes in both the intensity of competition across spatial replicates, and in resource availability through time, revealed changes in the targets of natural selection. Results provide empirical support for the long‐standing hypothesis that density‐dependent natural selection shapes the fitness surface of Greater Antilles anoles.     

Outdoor recreation causes effective habitat reduction in capercaillie Tetrao urogallus: a major threat for geographically restricted populations

Outdoor recreation inflicts a wide array of impacts on individual animals, many of them reflected in the avoidance of disturbed areas. The scale and spatial extent, however, at which wildlife populations are affected, are mostly unclear. Particularly in geographically isolated populations, where restricted habitat availability may preclude a relocation to undisturbed areas, effective habitat reduction may remain underestimated or even unnoticed, when animals stay in disturbed areas and only show small‐scale responses. Based on telemetry data, we investigated the spatial and seasonal effects of outdoor recreation – in relation to landscape and vegetation conditions – on western capercaillie Tetrao urogallus, considering two scales, home range and within‐home range habitat selection. We determined the distance‐thresholds up to which recreation infrastructures were avoided and estimated the extent of affected habitat for the isolated Black Forest (southwestern Germany) study population. While outdoor recreation did not affect home range selection, strong effects on habitat use within the home range were detected: distance to recreation infrastructure (hiking and cross‐country skiing trails, ski pistes) was the main determinant of habitat selection in winter; in summer, mountain bike trails and hiker's restaurants were avoided up to an average distance of 145 m (CI: 60–1092 m). Around winter‐infrastructure, relative avoidance was recorded up to 320 m (CI: 36–327 m), it was reduced, however, when dense understory provided visual cover. Of the entire population area, between 8–20% (summer) and 8–40% (winter) were affected by outdoor recreation, mainly in the high altitudes. Even without evident large‐scale shifts in species distribution, local‐scale avoidance of outdoor recreation can substantially contribute to effective habitat reduction. Based on our results we recommend a general reduction in recreation infrastructure density in key habitats, the establishment of undisturbed wildlife refuges with a diameter of at least 800 m, as well as enhancing visual protection by maintaining a strip of dense understory along trails.