Impact of prey occupancy and other ecological and anthropogenic factors on tiger distribution in Thailand's western forest complex

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Spatial variation in the density and vulnerability of preferred prey in the landscape shape patterns of Amur tiger habitat use

Theoretical and empirical research suggests that carnivore distributions are largely determined by prey availability. Availability depends not only on prey density but also on prey accessibility which is affected, in part, by the configuration of landscape attributes that make prey vulnerable to predation. Exactly how spatial variation in these processes shape patterns of carnivore habitat use at the home range scale remains poorly understood. We examined the influence of prey density (negative binomial resource selection function) and vulnerability (kill site resource selection function), mapped separately for each of three species of primary prey, on habitat use patterns within the home range for Amur tigers Panthera tigris altaica in Far East Russia over 20 winters. We developed spatially‐explicit mixed linear regression models to assess these patterns and found that models with parameters for specific primary prey were more robust than models with composite parameters for all primary prey species. This emphasizes the importance of evaluating predation dynamics at a species‐specific level. We also found that Amur tigers used habitat within the home range where red deer Cervus elaphus and wild boar Sus scrofa were dense. These two species were clearly preferred by tigers accounting for 72% (201 of the 278) of the tiger kills detected. The effect of red deer density however, was modulated by the vulnerability of red deer in the landscape. Amur tigers tended to establish their home ranges on habitat where red deer were most vulnerable to predation, but would use habitat where red deer were dense in the peripheral regions of their home ranges. This suggests that tigers may utilize two separate strategies for acquiring prey. As the configuration of resource patches within the home range influences carnivore survival and reproduction, our analysis has implications for tiger conservation that extend beyond our improved understanding of tiger‐prey ecology.     

Prey and tigers on the forgotten trail: high prey occupancy and tiger habitat use reveal the importance of the understudied Churia habitat of Nepal

Tigers are globally threatened and their conservation relies on intact habitat that supports key large prey. The Churia habitat is relatively unknown even though it occupies a significant portion of the forested landscape of the Terai Arc, which stretches over 1000 km in a narrow band across Nepal and India, parallel to the Himalayas. To address this lack of detailed information relevant to tiger conservation, we used sign surveys to estimate occupancy probability for 5 focal prey species of tigers (gaur, sambar, chital, wild pig, and barking deer), and assess tiger habitat use within 537 km² of the understudied Churia habitat in Chitwan National Park (CNP), Nepal. Multi-season occupancy models allowed us to make seasonal (winter vs. summer) inferences regarding changes in occupancy or habitat use based on covariates influencing occupancy and detection. We found that sambar had the largest spatial distribution occupying 431–437 km², while chital had the smallest at 100–158 km² across both seasons. The gaur population showed the most seasonal variation occupying from 413 to 318 km², suggesting their migration out of the Churia in summer and moving in during winter. Wild pigs showed the opposite trend occupying from 444 to 383 km²; suggesting moving into Churia in summer and out in winter. Barking deer were widespread in both seasons (329–349 km²). Tiger habitat use (\({{\hat{\varPsi } }}({\text{SE}})\)) was higher in winter 0.63 (0.11) than in summer 0.54 (0.21), but confidence intervals overlapped and area used was similar across seasons, 337 km² (winter) to 291 km² (summer). Available habitat, distribution of water sources, and human disturbance were the most common variables influencing spatial distribution of prey and habitat use of tigers at different spatial scales. Overall, we found high prey occupancy and tiger habitat use, suggesting the Churia is valuable habitat for ungulates and tigers. Given that this habitat accounts for 639 km² within CNP and 7642 km² across the entire Terai Arc, the Churia should no longer be neglected in global tiger conservation planning.     

Studying Large Mammals With Imperfect Detection: Status and Habitat Preferences of Wild Cattle and Large Carnivores in Eastern Cambodia

Studying large mammal species in tropical forests is a conservation challenge with species’ behavior and ecology often increasing the probability of non‐detection during surveys. Consequently, knowledge of the distribution, status, and natural history of many large mammal species in Southeast Asia is limited. I developed occupancy models from camera‐trapping data, thereby accounting for imperfect detection at sampling sites, to clarify the status and habitat requirements of four globally threatened or near threatened large mammals (banteng Bos javanicus, gaur Bos gaurus, dhole Cuon alpinus, and leopard Panthera pardus) in Mondulkiri Protected Forest, eastern Cambodia. Camera traps were operational for >3500 trap nights with 202 photographic encounters of the four study species. Model averaged occupancy estimates were between 5 percent (leopard) and 140 percent (gaur) higher than naive estimates (i.e., proportion of camera‐trap sites species recorded from) thus highlighting the importance of accounting for detectability during conservation surveys. I recommend the use of an occupancy framework when using camera‐trap data to study the status, ecology, and habitat preferences of poorly known and elusive species. The results highlight the importance of mixed deciduous and semi‐evergreen forest for wild cattle in eastern Cambodia and I emphasize that these habitats must be considered in conservation planning across the Lower Mekong Dry Forest Ecoregion.     

Molecular phylogeny of the tribe Bovini (Bovidae, Bovinae) and the taxonomic status of the Kouprey, Bos sauveli Urbain 1937

The kouprey is a very rare bovid species of the Indochinese peninsula, and no living specimen has been described for a long time, suggesting that it is possibly extinct. Its systematic position within the tribe Bovini remains confused since the analyses of morphological characters have led to several conflicting hypotheses. Some authors have also suggested that it could be a hybrid species produced by the crossing of the banteng with gaur, zebu, or water buffalo. Here we performed a molecular phylogeny of the tribe Bovini to determine the taxonomic status of the kouprey. DNA was extracted from the holotype specimen preserved in the MNHN collections. Phylogenetic analyses were carried out on a matrix including all the taxonomic diversity described in the tribe Bovini, and 2065 nucleotide characters, representing three different markers, i.e., the promotor of the lactoferrin and two mitochondrial genes (cytochrome b and subunit II of the cytochrome c oxidase). The results show that the kouprey belongs to the subtribe Bovina, and that three different clades can be evidenced into this group: the first includes the domestic ox, zebu, and European bison; the second incorporates the yak and American bison; and the third contains the kouprey, banteng and gaur. All hypotheses involving hybridization for the origin of the kouprey can be rejected, confirming that it is a real wild species. Molecular datings and biogeographic inferences suggest that the kouprey diverged from banteng and gaur during the Plio-Pleistocene of Asia. In addition, several molecular signatures were detected in the cytochrome b gene, permitting a molecular identification of the kouprey. We propose a conservation project based on a molecular taxonomy approach for tracking the kouprey in Indochina in order to determine whether some populations still survive in the wild.     

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.     

Diet and prey profiles of three sympatric large carnivores in Bandipur Tiger Reserve, India

We conducted a field study of diets of three sympatric large carnivores, the tiger Panthera tigris , the leopard Panthera pardus and the dhole Cuon alpinus in Bandipur Tiger Reserve, India, based on analyses of 381, 111 and 181 scats, respectively. The frequency of occurrence of prey items in scats was converted to relative biomass and number of prey consumed using regression equations based on earlier feeding trials. The results showed that although these predators kill ∼11–15 species of vertebrate prey, relatively abundant ungulate species provide 88–97% of biomass consumed by them. Although the dietary niche overlap among the three species was high (Pianka's index of 0.75–0.93), some specialized predation was observed. The largest ungulates, gaur Bos gaurus and sambar Cervus unicolor , provided 73% of biomass consumed by tigers, whereas medium-sized chital Axis axis and wild pig Sus scrofa formed 65 and 83% of the biomass intake of leopards and dholes, respectively. In terms of the relative numbers of prey animals killed by the three predators, chital, which is the most abundant prey species, dominated their diets (tiger=33%, leopard=39% and dhole=73%). The results of the study, in conjunction with earlier work, support the prediction that abundance of ungulate prey species, as well as their availability in different size classes, are both critical factors that facilitate sympatry among the three predators.     

Assessing habitat suitability for tiger in the fragmented Terai Arc Landscape of India and Nepal

Tiger Panthera tigris populations have declined dramatically in the Terai Arc Landscape (TAL; India and Nepal), and remaining populations are highly fragmented and endangered. As part of a research program to aid tiger management by identifying critical areas for conservation, we aimed to 1) identify the factors which affect the distribution of tigers in the TAL; 2) explore the role of spatial scale in habitat selection; 3) map potentially suitable habitats; and 4) assess the quality of potential corridors linking suitable habitats. We used an approach based on presence and pseudo‐absence data, combining ecological niche factor analysis and generalized linear models. We used an information‐theoretic approach to compare our data on tiger presence with different hypotheses on tiger habitat selection (i.e. protective habitat, prey species, human disturbance), and spatial scales. All hypotheses yielded models with high prediction accuracy (>79%). The most parsimonious model included variables characterizing habitat suitability of the 2 main prey species. More detailed assessment of potentially suitable areas using an extended source‐sink approach suggested that most of the habitats outside the protected areas were attractive sink‐like habitats (i.e. they suffered high levels of human disturbance in otherwise good habitats). Overall, 24% (ca 18 500 km²) of the study area was predicted as suitable (probability cut‐off p>0.5), approximately 7% of which is under protection. Our models showed that protecting the remaining concentrations of tigers requires focusing management efforts on specific areas outside the currently protected areas. These are characterized by good natural suitability; however, they suffer from a high level of human disturbance. Our models underscore the importance of minimizing human disturbances in these areas to avoid that they act as attractive sinks but act as corridors between existing subpopulations.     

Chronological distribution of the tiger Panthera tigris and the Asiatic lion Panthera leo persica in their common range in Asia

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Resolving a zoological mystery: the kouprey is a real species

The kouprey is a rare and enigmatic forest ox discovered by scientists in Cambodia only in 1937. Numerous morphological hypotheses have been proposed for the origin of the kouprey: that it is a species closely related to banteng and gaur, two other wild oxen of southeast Asia; a morphologically divergent species placed in a separate genus, named Novibos; a wild species linked to aurochs and domestic cattle; a vicariant population of banteng; a feral cattle; or a hybrid of banteng with either zebu cattle, gaur or water buffalo. In a recent paper, which gained a lot of media coverage, Galbreath et al. analysed mitochondrial DNA sequences and concluded that the kouprey never existed as a wild, natural species, and that it was a feral hybrid between banteng and zebu cattle. Here we analyse eight DNA markers-three mitochondrial regions and five nuclear fragments-representing an alignment of 4582 nucleotides for the holotype of the kouprey and all related species. Our results demonstrate that the kouprey is a real and naturally occurring species, and show that Cambodian populations of banteng acquired a mitochondrial genome of kouprey by natural introgressive hybridization during the Pleistocene epoch.     

亚洲虎种群恢复的机遇与挑战

虎(Panthera tigris)作为顶级捕食者, 对维持森林生态系统服务和结构完整性有着重要作用, 是研究和保护工作的旗舰物种。历史上, 虎曾广泛分布于亚洲大部分地区, 如今仅分布于南亚、东南亚和东北亚的破碎化栖息地, 各区域种群处于濒危或极度濒危状态。准确了解野生虎的种群状态和生态需求信息对于科学开展保护和恢复工作至关重要。本文通过综述近几十年的研究文献, 总结了野生虎种群现状和主要威胁因素, 评价了已有研究的重点与不足, 为未来亚洲虎种群的研究和保护提出了建议。目前与虎相关的研究主要集中在分布范围最广的孟加拉虎(P. t. tigris)和东北虎(P. t. altaica) 2个亚种, 而最急需关注的其他亚种仍研究不足。经过近十几年的努力和保护投入, 目前野生虎种群数量已从2010年的大约3,200只恢复到现在约4,500只, 但在越南、柬埔寨和老挝3个国家已经灭绝。虎面临的主要威胁包括持续的栖息地破坏和隔离、猎物缺乏、近交衰退、人虎冲突、贸易与盗猎和疾病威胁等。未来的研究和保护工作需要加强种群和栖息地连通性恢复、个体重引入、疾病管控以及加强跨境合作和反盗猎等。     

福建梅花山圈养华南虎遗传状况分析

华南虎是世界上密切关注的旗舰物种,在过去的10 年间没有发现野生华南虎存在的证据,因此它是极度濒危的虎亚种。福建梅花山圈养华南虎群体是整个圈养华南虎群体的重要组成部分,拥有 12 只华南虎。基于组合长度为 3934 bp 的线粒体序列分析发现梅花山圈养华南虎拥有 3 种线粒体单倍型; 而基于 20 个微卫星位点基因型分析显示梅花山圈养华南虎一共有71 个等位基因,平均等位基因数是3. 55,等位基因丰度的平均值是3. 32,平均期望杂合度和多态信息含量( PIC) 分别为0. 513 和0. 445。这些提示梅花山华南虎圈养群体维持着较高的遗传多样性。     

Effects of human‐induced prey depletion on large carnivores in protected areas: Lessons from modeling tiger populations in stylized spatial scenarios

Prey depletion is a major threat to the conservation of large carnivore species globally. However, at the policy‐relevant scale of protected areas, we know little about how the spatial distribution of prey depletion affects carnivore space use and population persistence. We developed a spatially explicit, agent‐based model to investigate the effects of different human‐induced prey depletion experiments on the globally endangered tiger (Panthera tigris) in isolated protected areas—a situation that prevails throughout the tiger's range. Specifically, we generated 120 experiments that varied the spatial extent and intensity of prey depletion across a stylized (circle) landscape (1,000 km²) and Nepal's Chitwan National Park (~1,239 km²). Experiments that created more spatially homogenous prey distributions (i.e., less prey removed per cell but over larger areas) resulted in larger tiger territories and smaller population sizes over time. Counterintuitively, we found that depleting prey along the edge of Chitwan National Park, while decreasing tiger numbers overall, also decreased female competition for those areas, leading to lower rates of female starvation. Overall our results suggest that subtle differences in the spatial distributions of prey densities created by various human activities, such as natural resource‐use patterns, urban growth and infrastructure development, or conservation spatial zoning might have unintended, detrimental effects on carnivore populations. Our model is a useful planning tool as it incorporates information on animal behavioral ecology, resource spatial distribution, and the drivers of change to those resources, such as human activities.     

亚洲同域分布虎和豹竞争与共存关系概述

虎(Panthera tigris)和豹(P. pardus)作为食物链的顶级捕食者对维持生态系统结构与功能稳定性起到重要作用。强烈的人为干扰已导致亚洲虎和豹种群(以下简称虎豹)分布呈现破碎化状态, 探究二者之间的相互作用对缓解人类威胁和濒危物种保护具有重要意义。尽管虎豹竞争和共存研究已持续几十年, 但多局限于区域性种群, 缺乏系统性研究。本研究梳理了1976-2021年间36篇涉及虎豹竞争性相互作用研究的文献, 涵盖中国、俄罗斯、泰国、马来西亚、印度、尼泊尔、不丹共7个国家26个区域, 系统阐述了虎豹目前在亚洲的主要分布以及它们之间的干涉型和资源利用型竞争的主要表现形式, 并分析了猎物与干扰因素对虎豹竞争与共存的影响。本研究提出了5个可能影响种间作用的猎物构成情况, 强调了不同大小有蹄类猎物的丰富度和多度以及人为干扰的空间分布对虎豹区域竞争和共存的调控作用。现有研究显示虎豹的空间利用取决于本地的生境、猎物和干扰等因素。豹相对于虎具有更强的行为可塑性, 虎豹之间在空间、时间和营养生态位等维度权衡生态机会(如容易捕获的猎物)与冲突风险(人或竞争者带来的风险)来促进共存。目前虎豹相互作用研究存在严重的亚种和区域不平衡性, 未来研究重点应关注各种生物和非生物类因素对虎豹种间竞争类型和作用强度的调控方式以及调控的生态阈值效应。     

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.     

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）。     

Responses of tiger (<Emphasis Type="Italic">Panthera tigris</Emphasis>) and their prey to removal of anthropogenic influences in Rajaji National Park,India

Presence of human settlements in most protected areas has forced tigers (Panthera tigris) to share space with humans. Creation of inviolate space for tigers in areas with high human densities is often daunting and requires hard political sacrifices. We conducted this study from 2004 to 2007 in the Chilla range of Rajaji National Park, along the northwestern portion of the Terai-Arc Landscape in the Indian subcontinent. Our objective was to document the recovery of prey and tiger populations following the resettlement of 193 gujjar (pastoralists with large buffalo holdings) families. We used distance sampling to estimate density of wild ungulate prey and camera traps to estimate tiger density. The study area supported ∼66 ungulates/km², with chital (Axis axis) and sambar (Cervus unicolor) contributing >91%. While prey densities did not vary across 3 years, an increase in proportion of chital fawns was observed following the near complete removal of livestock. We also documented an increase in the density of tigers (from three to five tigers per 100 km²), probably due to immigrating tigers from nearby Corbett Tiger Reserve. A high turnover of individual tigers was observed during the study. With photographic evidence of breeding tigers in Chilla range, we believe that this area could serve as a source population from where tigers can colonize adjoining forests across River Ganga. It is therefore concluded that securing the connectivity between forests on the east and west bank of Ganga through the tenuous Chilla-Motichur corridor assumes significance for long-term persistence of tigers within this landscape.     

Tiger straying incidents in Indian Sundarban: statistical analysis of case studies as well as depredation caused by conflict

The Sundarban of India and Bangladesh is the only mangrove reserve forest in the world inhabited by the tiger (Panthera tigris). Tigers in the Sundarban mangrove are widely known for frequently straying into the surrounding reclaimed areas. Data collected from household village survey and documents of the Forest Department show that tiger straying incidents happen throughout the year, but most of them occurred during 3 months (Dec–Feb) of the winter season (42%) followed by 3 months (July–Sept) of the monsoon season (31%). 84.22% of cases have been reported from 21 villages of five affected blocks of Sundarban. In most cases, tigers resorted to cattle lifting or poultry feeding. Only in 8.9% of the cases were human beings attacked or killed. Majority of the straying tigers (68.46%) were male. In most cases (78.9%) strayed tigers were aged and 22% of these were partly injured. 96.05% straying occurs during night. This study also aims at exploring the causes of frequent straying, livestock and human casualties as a result of conflict and retaliation killing of tigers. Straying frequency is correlated negatively with the width of the creeks or rivers in the village side and no relationship is identified with the area of the forest block as well as natural prey abundance. Overall, improved nylon fencing, increased patroling, establishment of the Forest Protection Committee (FPC) and the Eco Development Committee (EDC) are not associated with reduction of straying frequency as well as livestock losses to tiger straying.     

Life‐history traits of gaur Bos gaurus: a first analysis

In this first detailed analysis of gaur Bos gaurus life‐history traits, data were collected from a 20‐month field study in South India and from captive gaur populations. Mean age of females at first parturition was 3 years; females remained fertile beyond the age of 15 years. Adult females were three times more abundant than adult males in the wild; survival of females was greater than males beyond three years of age. Life span of both sexes has not exceeded 24 years in captivity. Gaur life‐history traits are similar to those of other similar‐sized Bovini species.