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。     

黑龙江省完达山东部林区东北虎猎物生物量

研究一个地区猎物种群生物量能否满足捕食动物种群数量的需求,这对于了解濒危大型食肉动物是否受到来自于食物缺乏的威胁和制定相应的保护措施极其重要。为了掌握黑龙江省完达山东部林区东北虎食物需求与猎物生物量之间的关系,于2008年冬季至2009早春积雪覆盖期采用随机布设样线,通过收集有蹄类动物在雪地上留下的足迹等活动的方法,在东方红林业局和迎春林业局管辖境内3 692.06 km²的区域布设大样方48个,并在大样方里共布设样线240条开展有蹄类动物种群数量调查,确定东北虎猎物生物量。调查结果表明:研究地区野猪(成体502 606只,亚成体209 210只)、马鹿(成体331 357只,亚成体67 72只)和狍子(成体810 815只,亚成体202 203只)的生物量分别为74 767.50 87 825.00 kg、79 744.50 85 984.50 kg 和 31 337.00 31 525.50 kg,3种有蹄类动物生物量共计1 85 849.00 205 335.00 kg。研究地区猎物总生物量为209 619.89 231 598.24 kg。如果按8%的生物提供给东北虎,3种主要猎物生物量可满足5.22 6.92只东北虎个体的食物需求,研究地区猎物总生物量则可满足5.89 7.81只东北虎个体的食物需求。此外,对足迹遇见率与抽样强度、抽样强度与足迹遇见率的均值标准误差之间关系的分析表明,在完达山东部林区布设120条样线(抽样距离600 km)、150条样线(抽样距离750 km)和115条样线(抽样距离675 km)能满足野猪、马鹿、狍子种群数量调查准确性的最低需求。     

吉林省东北虎的调查

经过8年的调查研究,确定了东北虎在吉林省的分布区有3处,数量为7－9只。即大龙岭分布区3－5只,哈尔巴岭分布区1只,张广才岭分布区3只。东北虎的适宜生境为海拔800－1100m的中低山：人口压力在15人／km^2以下,被捕食动物野猪、狍、马鹿的密度在2．5只／km^2以上。林型为以柞木为主的阔叶混交林,林龄为中成林。     

Prey Animals of Tiger (<Emphasis Type="Italic">Panthera tigris tigris</Emphasis>) in Dudhwa Landscape,Terai Region,North India

Livestock depredation by carnivores cause substantial human carnivore conflict and subsequently decreased support for carnivore conservation. Thus, understanding carnivore diet with respect to wild prey availability has major implications to determine the reasons behind livestock depredation. A study was conducted to investigate food habits and prey use of tiger at four study sites (Dudhwa National Park, Kishanpur Wildlife Sanctuary, Pilibhit Forest Division and Katerniaghat Wildlife Sanctuary) in Dudhwa landscape, Terai Region, North India for further understanding of prey–predator relationship and partial impact of wild prey availability on livestock depredation by tiger through scat analysis. Scat analysis shows that the tigers depend mostly on medium sized prey throughout the study area (74.11, 73.58, 71.79, 47.62%). In Dudhwa National Park and Kishanpur Wildlife Sanctuary, predation was attempted subsequently on wider prey variety of eleven and nine different available prey species where livestock depredation were only 3.77 and 5.36% respectively. While, in absence of wider prey variety, large sized livestock (21.91, 16.55%) and nilgai (24.41, 5.57%) contributed much higher in tiger diet in Pilibhit Forest Division and Katerniaghat Wildlife Sanctuary respectively. Our study suggested that availability of prey variety has an important role in reduced livestock depredation. Medium sized preys were mostly contributing in tiger diet and seems to be a significant parameter for sustaining tiger population where abundance of large sized prey is rare. Conservation of medium sized preys is important but along with natural restoration of the population of large sized prey species like sambar and swamp deer is essential in order to reduce livestock depredation.     

Effects of human–carnivore conflict on tiger (Panthera tigris) and prey populations in Lao PDR

Unique to South-east Asia, Lao People's Democratic Republic contains extensive habitat for tigers and their prey within a multiple-use protected area system covering 13% of the country. Although human population density is the lowest in the region, the impact of human occurrence in protected areas on tiger Panthera tigris and prey populations was unknown. We examined the effects of human–carnivore conflict on tiger and prey abundance and distribution in the Nam Et-Phou Louey National Protected Area on the Lao–Vietnam border. We conducted intensive camera-trap sampling of large carnivores and prey at varying levels of human population and monitored carnivore depredation of livestock across the protected area. The relative abundance of large ungulates was low throughout whereas that of small prey was significantly higher where human density was lower. The estimated tiger density for the sample area ranged from 0.2 to 0.7 per 100 km². Tiger abundance was significantly lower where human population and disturbance were greater. Three factors, commercial poaching associated with livestock grazing followed by prey depletion and competition between large carnivores, are likely responsible for tiger abundance and distribution. Maintaining tigers in the country's protected areas will be dependent on the spatial separation of large carnivores and humans by modifying livestock husbandry practices and enforcing zoning.     

Tiger and leopard co-occurrence: intraguild interactions in response to human and livestock disturbance

Intraguild interactions have important implications for carnivore demography and conservation. Differences in how predators respond to different forms of disturbance might alter their interaction patterns. We sought to understand how human and livestock disturbance impact co-occurrence of sympatric large carnivores such as tiger (Panthera tigris) and leopard (P. pardus) and thereby mediate the intraguild interaction pattern to enable coexistence of the species in a human-dominated landscape. We surveyed 361 locations in Chitwan National Park, Nepal, to examine how prey abundance and disturbance factors such as human and livestock presence might influence habitat use by tigers and leopards independently and when co-occurring. Single-species single-season models and two-species single-season models were developed to examine hypotheses on unconditional detection and occupancy and species interaction respectively. Pervasive human use of the park had negative impacts on tiger occupancy while the abundance of prey had a positive influence. Despite significant prey overlap between tigers and leopards, none of the native prey species predicted leopard habitat occupancy. However, habitats used extensively by livestock were also used by leopards. Further, we found strong evidence of intraguild competition. For instance tiger occupancy was higher in prey-rich areas and leopard occupancy was low in the sites where tigers were present. These findings, and a species interaction factor of < 1 clearly indicate that leopards avoid tigers, but their use of areas of disturbance enables them to persist in fringe habitats. We provide empirical evidence of how intraguild interaction may result in habitat segregation between competing carnivores, while also showing that human and livestock use of the landscape create disturbance patterns that facilitate co-occurrence of the predators. Thus, because large carnivores compete, some disturbance may mediate coexistence in small protected areas. Understanding such interactions can help address important conservation challenges associated with maintaining diverse carnivore communities in small or disturbed landscapes.     

Weights of gaur (Bos gaurus) and banteng (Bos javanicus) killed by tigers in Thailand

The primary prey of tigers across much of South‐East Asia has been depleted, reducing the ability of already limited habitat to support tigers. To better understand the extent to which two of the largest prey species, gaur (Bos gaurus) and banteng (Bos javanicus), contribute to the tiger's diet, we estimated the average size of these species killed by tigers. This information is needed to more accurately calculate biomass of these species in the tiger's diet and to devise strategies to increase tiger carrying capacity where habitat is fragmented and limited in west‐central Thailand. We used temporally clumped locations of 24 satellite radio‐collared tigers to identify their kill sites and obtained mandibles from 82 gaur and 79 banteng. Kills were aged by teeth eruption sequence, sectioning the M1 molar and counting cementum annuli. Of all gaur killed, 45.2% were adults; of all banteng killed, 55.7% were adults. The average weight of banteng killed was 423.9 kg, which was similar to the 397.9 kg average weight for gaur. The mean weight of both prey species is 3.5–4.5 times greater than the predicted 1:1 preferred prey to predator ratio. In the absence of medium‐sized prey, killing these larger animals may be especially critical for female tigers provisioning nearly independent young when male offspring are already larger than the mother. This is the first study to present data on the average weights of gaur and banteng killed in South‐East Asia, and these results suggest that these are key prey species to target in tiger prey recovery efforts.     

Competition versus opportunism: Diet and trophic niche relationship of two sympatric apex carnivores in a tropical forest

There is a little understanding of how apex carnivores partition their diet to coexist. We studied food habit and trophic niche overlap of two apex carnivores, tiger and leopard in the tropical forest of Similipal tiger reserve, eastern India. We used line transect and scat analysis method, to estimate the prey availability and determine the diet and prey selection of two apex carnivores. Tigers consumed mostly large and medium-sized prey, whereas leopards mostly consumed medium and small-sized prey. Both carnivores were not random in their consumption of prey, sambar and wild pigs were selectively consumed by tigers, whereas leopards selectively consumed wild pigs, barking deer and mouse deer. Dietary overlap between two carnivore species was moderate (Pianka's niche overlap index: 0.55), and trophic niche breadth (Levin's standardized niche breadth) of the leopard (0.52) was greater than that of the tiger (0.37). Overall, tigers exhibited specialized feeding habits, whereas leopards showed generalist feeding habits. Our study highlights the opportunistic nature of leopards and probably a reason for the species successfully coexist with tigers.     

张广才岭南部东北虎猎物种群冬季营养容纳量研究

猎物匮乏是影响东北虎（Panthera tigris altaica）种群恢复的关键因素之一。容纳量研究是开展东北虎猎物恢复工作的必要前提。通过Maxent模型、聚类分析和训练随机树分类等方法,结合调查数据,预测了吉林省张广才岭南部黄泥河林业局东北虎主要猎物的适宜栖息地空间分布,解译了植被类型,在此基础上基于不同植被类型动物可采食部分代谢能、不同生境等级食物可利用率、马鹿（Cervus elaphus）和狍（Capreolus capreolus）生境等级重叠情况以及动物冬季能量需求,分析了东北虎猎物的冬季营养容纳量。结果表明：黄泥河林业局狍、野猪（Sus scrofa）和马鹿的适宜栖息地分别占研究区域总面积的52.8%、40.7%和25.4%;从猎物获取能量来看,以山杨（Populus davidiana）、桦树（Betula）、核桃楸（Juglans mandshurica）为主的植被类型是马鹿、狍可获得能量较多的生境,以蒙古栎（Quercus mongolica）、核桃楸为主的植被类型是野猪可获得能量较多的生境。东北虎猎物种群的综合冬季营养容纳量为574只马鹿（0.29只/km²）,7016只狍（3.54只/km²）,4785只野猪（2.38只/km²）。     

Landscape‐scale accessibility of livestock to tigers: implications of spatial grain for modeling predation risk to mitigate human–carnivore conflict

Innovative conservation tools are greatly needed to reduce livelihood losses and wildlife declines resulting from human–carnivore conflict. Spatial risk modeling is an emerging method for assessing the spatial patterns of predator–prey interactions, with applications for mitigating carnivore attacks on livestock. Large carnivores that ambush prey attack and kill over small areas, requiring models at fine spatial grains to predict livestock depredation hot spots. To detect the best resolution for predicting where carnivores access livestock, we examined the spatial attributes associated with livestock killed by tigers in Kanha Tiger Reserve, India, using risk models generated at 20, 100, and 200‐m spatial grains. We analyzed land‐use, human presence, and vegetation structure variables at 138 kill sites and 439 random sites to identify key landscape attributes where livestock were vulnerable to tigers. Land‐use and human presence variables contributed strongly to predation risk models, with most variables showing high relative importance (≥0.85) at all spatial grains. The risk of a tiger killing livestock increased near dense forests and near the boundary of the park core zone where human presence is restricted. Risk was nonlinearly related to human infrastructure and open vegetation, with the greatest risk occurring 1.2 km from roads, 1.1 km from villages, and 8.0 km from scrubland. Kill sites were characterized by denser, patchier, and more complex vegetation with lower visibility than random sites. Risk maps revealed high‐risk hot spots inside of the core zone boundary and in several patches in the human‐dominated buffer zone. Validation against known kills revealed predictive accuracy for only the 20 m model, the resolution best representing the kill stage of hunting for large carnivores that ambush prey, like the tiger. Results demonstrate that risk models developed at fine spatial grains can offer accurate guidance on landscape attributes livestock should avoid to minimize human–carnivore conflict.     

黑龙江省东完达山地区东北虎猎物种群现状及动态趋势

猎物种群丰度是限制虎分布和数量的关键因子,因此猎物种群密度监测和估算是虎保护的重要内容之一。应用采用大样方法,地理信息系统技术和多元统计分析,研究了黑龙江东完达山东部地区东北虎猎物种群(马鹿、狍子和野猪)现状及动态变化趋势。结果表明:研究地区马鹿的种群平均密度为(0.2010±0.0270)只/km²、狍子的平均种群密度为(0.4980±0.0436)只/km²、野猪的平均种群密度为(0.3423±0.0275)只/km²。单因素方差分析表明,在相同生境下,3种有蹄类密度在在阔叶混交林中和杂木林中差异极为显著;不同的生境,3种猎物的猎物的密度也存在着显著差异。相关分析表明马鹿密度和野猪密度程正相关,而马鹿密度和狍子密度、狍子密度和野猪密度则不相关。 同1989年该地区东北虎猎物种群相比:1989-2002年的13 a时间内马鹿的年平均递减率为13.48%、狍子的年平均递减率为12.69%、野猪的年平均递减率为1.89%。     

Prey selection and dietary response by wolves in a high-density multi-species ungulate community

Studies on predation by the wolf (Canis lupus) have often reported contradictory results about the role of prey density and vulnerability on wolf prey use. We investigated dietary response and prey selection by wolves in a high-density and multi-species ungulate community, analysing scats collected over a period of 11 years in the Casentinesi Forests, Italy. The second most abundant species, wild boar (Sus scrofa), was found to be the main wolf prey, and we did not observe any dietary response of wolves to variations in the density of either primary or secondary prey species. Selection patterns were uniform throughout the study period. Wolves strongly selected for wild boar piglets, while roe deer (Capreolus capreolus) fawns and adults, red deer (Cervus elaphus) adults and fallow deer (Dama dama) adults were avoided. Wolf preference for wild boar was inversely density dependent. Within each species, juveniles were preferred to adults. Medium-sized, young individuals of both wild boar and roe deer were optimal prey, although with different selection patterns related to the different anti-predator strategies adopted by each prey species. The results of this study suggest that in productive ecosystems with high density and high renewal rates of prey, selection patterns by wolves are determined by prey vulnerability, which is connected to prey age and body size. The different patterns of wild boar versus cervids use by wolf across Europe seems to be related to their relative abundances, while the strong selection of wild boar in Italian Apennines with respect to the more frequent avoidance in central-eastern Europe is better explained by higher piglet productivity and smaller body size of adults boar in Mediterranean temperate forests.     

Effects of environmental and anthropogenic drivers on Amur tiger distribution in northeastern China

We examined environmental and anthropogenic factors drive range loss in large mammals, using presence data of Amur tigers opportunistically collected between 2000 and 2012, and anthropogenic and environmental variables to model the distribution of the Amur tiger in northeastern China. Our results suggested that population distribution models of different subregions showed different habitat factors determining tiger population distribution patterns. Where farmland cover was over 50 km² per pixel (196 km²), distance was within 15 km to the railway in Changbaishan and road density (length per pixel) increased in Wandashan, the relative probability of Amur tiger occurrence exhibited monotonic avoidance responses; however, where distance was within 150 km of the Sino-Russia border, the occurrence probability of Amur tiger was relatively high. We analyzed the avoidance or preference responses of Amur tiger distribution to elevation, snow depth and Viewshed. Furthermore, different subregional models detected a variety of spatial autocorrelation distances due to different population clustering patterns. We found that spatial models significantly improved model fits for non-spatial models and made more robust habitat suitability predications than that of non-spatial models. Consequently, these findings provide useful guidance for habitat conservation and management.     

中国野生东北虎数量监测方法有效性评估

虎数量监测是虎保护的核心内容之一。野生虎现存数量少、领域宽广,加之习性机警,很难对其数量和种群变化趋势做出准确的评估。合适的虎数量监测方法随着监测目标、监测尺度、虎密度、猎物密度、气候及其它环境因素的变化而不同。2002—2011年,用东北虎信息收集网络法,样线调查法,猎物生物量和捕食者关系法,对东北虎数量进行监测。(1)用老虎信息收集网络法研究2006年完达山东部地区东北虎的种群现状,结果显示东完达山地区2006年东北虎数量为6—9只,由1只成年雄虎,2—3只成年雌虎,2—4只亚成体虎和1只小于1岁的幼体虎组成;(2)用猎物生物量和捕食者关系法得到东完达山地区2002年东北虎的密度为0.356只/100 km2,能容纳22—27只东北虎;(3)用样线法在黑龙江的老爷岭南部和吉林省大龙岭北部面积1735.99 km2的区域内设置样线64条,总长609 km,没有发现东北虎足迹链。样线调查的结果表明,在2011年2—3月该调查区域东北虎的数量为0只。监测结果表明,用猎物生物量和捕食者关系得到东北虎数量远远超过现实数量,人们对有蹄类的盗猎和猎套对老虎的伤害可能是其主要原因;样线法调查得出的结果低于现实种群,主要原因是老虎数量极低和调查者对野生虎行为学了解甚少,较难在野外有效的发现虎信息;且样线法监测仅应用于当东北虎以一定的密度(即有定居虎)存在的情况下(多数监测样线能发现虎信息)。虽然和样线法一样存在着诸如专家估计密度和真实密度之间的关系,老虎足迹数量和老虎真实密度间关系不确定,保守估计等内在缺点,在目前中国东北地区野生东北虎种群密度极低,且多是穿越于中俄边境地区的游荡个体的现状下,信息收集网络法是一种高效,可行东北虎监测方法。因此,建议建立更广泛的监测信息收集网络,培训监测人员,严格执行信息收集程序,减少专家估计误差以完善此监测方法。此外,其他监测方法,如占有法、基于标志重捕远红外照相法、粪便DNA法、足迹数码信息法、警犬法等,应根据各种方法的理论前提、误差来源、适用范围和老虎是否定居及密度等具体情况有选择地加以应用,且有些方法可能成为未来中国野生东北虎种群的有效监测工具。     

The basic causes of death in hoofed mammals in the ussuri reserve and in the adjacent territory

The basic causes of death in hoofed mammals in the Urruri reserve and in the adjacent territory during the period from 1974 to 2011 are hunting and poaching, tiger predation, and exhaustion. The sika deer has accounted for the largest number of the established deaths since the 1990s, because this species has become dominant in the fauna of hoofed animals in the reserve as a result of acclimatization. Before the 1980s, the main prey for tigers were the red deer (52%) and wild boar (44%): the share of the sika deer in the prey was 4%. During the most recent 10 years, the share of the sika deer, wild boar, and red deer in the tiger-prey trophic link at separate stations of the reserve has been 68%, 20.6%, and 6.9%, respectively.     

Spatial Partitioning of Predation Risk in a Multiple Predator-Multiple Prey System

ABSTRACT Minimizing risk of predation from multiple predators can be difficult, particularly when the risk effects of one predator species may influence vulnerability to a second predator species. We decomposed spatial risk of predation in a 2-predator, 2-prey system into relative risk of encounter and, given an encounter, conditional relative risk of being killed. Then, we generated spatially explicit functions of total risk of predation for each prey species (elk [Cervus elaphus] and mule deer [Odocoileus hemionus]) by combining risks of encounter and kill. For both mule deer and elk, topographic and vegetation type effects, along with resource selection by their primary predator (cougars [Puma concolor] and wolves [Canis lupus], respectively), strongly influenced risk of encounter. Following an encounter, topographic and vegetation type effects altered the risk of predation for both ungulates. For mule deer, risk of direct predation was largely a function of cougar resource selection. However, for elk, risk of direct predation was not only a function of wolf occurrence, but also of habitat attributes that increased elk vulnerability to predation following an encounter. Our analysis of stage-based (i.e., encounter and kill) predation indicates that the risk effect of elk shifting to structurally complex habitat may ameliorate risk of direct predation by wolves but exacerbate risk of direct predation by cougars. Information on spatiotemporal patterns of predation will be become increasingly important as state agencies in the western United States face pressure to integrate predator and prey management.     

Estimating the Population Size and Genetic Diversity of Amur Tigers in Northeast China

Over the past century, the endangered Amur tiger (Panthera tigris altaica) has experienced a severe contraction in demography and geographic range because of habitat loss, poaching, and prey depletion. In its historical home in Northeast China, there appears to be a single tiger population that includes tigers in Southwest Primorye and Northeast China; however, the current demographic status of this population is uncertain. Information on the abundance, distribution and genetic diversity of this population for assessing the efficacy of conservation interventions are scarce. We used noninvasive genetic detection data from scats, capture-recapture models and an accumulation curve method to estimate the abundance of Amur tigers in Northeast China. We identified 11 individual tigers (6 females and 5 males) using 10 microsatellite loci in three nature reserves between April 2013 and May 2015. These tigers are confined primarily to a Hunchun Nature Reserve along the border with Russia, with an estimated population abundance of 9–11 tigers during the winter of 2014–2015. They showed a low level of genetic diversity. The mean number of alleles per locus was 2.60 and expected and observed heterozygosity were 0.42 and 0.49, respectively. We also documented long-distance dispersal (~270 km) of a male Amur tiger to Huangnihe Nature Reserve from the border, suggesting that the expansion of neighboring Russian populations may eventually help sustain Chinese populations. However, the small and isolated population recorded by this study demonstrate that there is an urgent need for more intensive regional management to create a tiger-permeable landscape and increased genetic connectivity with other populations.     

西双版纳尚勇自然保护区野生印支虎及其三种主要有蹄类猎物种群现状调查

中国野生印支虎及其猎物种群状况的野外实地研究一直处于空白。本研究使用足迹鉴别法、粪堆计数法,首次对西双版纳尚勇自然保护区野生印支虎种群数量现状及该区域内的虎猎物种群状况进行了调查研究。结果显示:2004 ～ 2009 年间,确认西双版纳保护区存在3 只成年印支虎个体(2 雌1 雄),西双版纳尚勇保护区拥有比较丰富的有蹄类种群,其中虎的主要猎物:水鹿平均密度为7.63 (7.40 ～ 9.23)只/ km² ;赤麂平均密度为17. 39 (11.33 ～24.94)只/ km² ,野猪平均密度为10.26 (7.69 ～ 14.51) 只/ km² ,该区域虎猎物生物量为1 715. 74 kg/ km² 。本研究还探讨了该区域印支虎种群的保护前景以及中国境内开展虎种群调查的适用办法等。     

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.     

The landscape of fear as an emergent property of heterogeneity: Contrasting patterns of predation risk in grassland ecosystems

The likelihood of encountering a predator influences prey behavior and spatial distribution such that non‐consumptive effects can outweigh the influence of direct predation. Prey species are thought to filter information on perceived predator encounter rates in physical landscapes into a landscape of fear defined by spatially explicit heterogeneity in predation risk. The presence of multiple predators using different hunting strategies further complicates navigation through a landscape of fear and potentially exposes prey to greater risk of predation. The juxtaposition of land cover types likely influences overlap in occurrence of different predators, suggesting that attributes of a landscape of fear result from complexity in the physical landscape. Woody encroachment in grasslands furnishes an example of increasing complexity with the potential to influence predator distributions. We examined the role of vegetation structure on the distribution of two avian predators, Red‐tailed Hawk (Buteo jamaicensis) and Northern Harrier (Circus cyaneus), and the vulnerability of a frequent prey species of those predators, Northern Bobwhite (Colinus virginianus). We mapped occurrences of the raptors and kill locations of Northern Bobwhite to examine spatial vulnerability patterns in relation to landscape complexity. We use an offset model to examine spatially explicit habitat use patterns of these predators in the Southern Great Plains of the United States, and monitored vulnerability patterns of their prey species based on kill locations collected during radio telemetry monitoring. Both predator density and predation‐specific mortality of Northern Bobwhite increased with vegetation complexity generated by fine‐scale interspersion of grassland and woodland. Predation pressure was lower in more homogeneous landscapes where overlap of the two predators was less frequent. Predator overlap created areas of high risk for Northern Bobwhite amounting to 32% of the land area where landscape complexity was high and 7% where complexity was lower. Our study emphasizes the need to evaluate the role of landscape structure on predation dynamics and reveals another threat from woody encroachment in grasslands.