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。     

Ranging,Activity and Habitat Use by Tigers in the Mangrove Forests of the Sundarban

The Sundarban of India and Bangladesh (about 6000 km²) are the only mangrove forests inhabited by a sizeable population of tigers. The adjoining area also supports one of the highest human densities and experiences severe human-tiger conflicts. We used GPS-Satellite and VHF radio-collars on 6 (3 males and 3 female) tigers to study their ranging patterns and habitat preference. The average home range (95% Fixed Kernel) for resident females was 56.4 (SE 5.69) and for males it was 110 (SE 49) km². Tigers crossed an average of 5 water channels > 30 meters per day with a mean width of 54 meters, whereas channels larger than 400 meters were rarely crossed. Tigers spent over 58% of their time within Phoenix habitat but compositional analysis showed a habitat preference of the order Avicennia-Sonneratia > Phoenix > Ceriops > Barren > Water. Average daily distance moved was 4.6 km (range 0.1–23). Activity of tigers peaked between 05:00 hours and 10:00 hours showing some overlap with human activity. Territory boundaries were demarcated by large channels which tigers intensively patrolled. Extra caution should be taken while fishing or honey collection during early morning in Avicennia-Sonneratia and Phoenix habitat types along wide channels to reduce human-tiger conflict. Considering home-range core areas as exclusive, tiger density was estimated at 4.6 (SE range 3.6 to 6.7) tigers/100 km² giving a total population of 76 (SE range 59–110) tigers in the Indian Sundarban. Reluctance of tigers to cross wide water channels combined with increasing commercial boat traffic and sea level rise due to climate change pose a real threat of fragmenting the Sundarban tiger population.     

The year of the tiger and the year of tiger genomes!

Tigers are endangered apex predators. They typify endangered species because they are elusive, rare, and face numerous threats across their range. Tigers also symbolize conservation. However, it is very difficult to study tigers because of their stated nature. Also, tiger conservation is a geopolitically sensitive topic, making it difficult to use the studies to propose evidence-based management that allows their recovery, especially in the context of conservation genetics. Zhang et al. (Mol. Ecol. Resour., 2022) have created very valuable and rare resources to aid the community in conserving tigers. First, they present chromosome level genome assemblies of a South China tiger and an Amur tiger. Second, they present whole genome sequences of 16 captive South China tigers. Additionally, by using the assemblies they model the demographic history of these populations, estimate inbreeding and the potential threats they face in captivity. This approach is particularly important because genetic management is now the only remaining option for South China tigers, because they are extinct in the wild. In other words, captive individuals are our only hope for some day restoring the wild populations of South China tigers.     

Recent Evolutionary History of Tigers Highlights Contrasting Roles of Genetic Drift and Selection

Species conservation can be improved by knowledge of evolutionary and genetic history. Tigers are among the most charismatic of endangered species and garner significant conservation attention. However, their evolutionary history and genomic variation remain poorly known, especially for Indian tigers. With 70% of the world’s wild tigers living in India, such knowledge is critical. We re-sequenced 65 individual tiger genomes representing most extant subspecies with a specific focus on tigers from India. As suggested by earlier studies, we found strong genetic differentiation between the putative tiger subspecies. Despite high total genomic diversity in India, individual tigers host longer runs of homozygosity, potentially suggesting recent inbreeding or founding events, possibly due to small and fragmented protected areas. We suggest the impacts of ongoing connectivity loss on inbreeding and persistence of Indian tigers be closely monitored. Surprisingly, demographic models suggest recent divergence (within the last 20,000 years) between subspecies and strong population bottlenecks. Amur tiger genomes revealed the strongest signals of selection related to metabolic adaptation to cold, whereas Sumatran tigers show evidence of weak selection for genes involved in body size regulation. We recommend detailed investigation of local adaptation in Amur and Sumatran tigers prior to initiating genetic rescue.     

Forest corridors maintain historical gene flow in a tiger metapopulation in the highlands of central India

Understanding the patterns of gene flow of an endangered species metapopulation occupying a fragmented habitat is crucial for landscape-level conservation planning and devising effective conservation strategies. Tigers (Panthera tigris) are globally endangered and their populations are highly fragmented and exist in a few isolated metapopulations across their range. We used multi-locus genotypic data from 273 individual tigers (Panthera tigris tigris) from four tiger populations of the Satpura–Maikal landscape of central India to determine whether the corridors in this landscape are functional. This 45 000 km² landscape contains 17% of India''s tiger population and 12% of its tiger habitat. We applied Bayesian and coalescent-based analyses to estimate contemporary and historical gene flow among these populations and to infer their evolutionary history. We found that the tiger metapopulation in central India has high rates of historical and contemporary gene flow. The tests for population history reveal that tigers populated central India about 10 000 years ago. Their population subdivision began about 1000 years ago and accelerated about 200 years ago owing to habitat fragmentation, leading to four spatially separated populations. These four populations have been in migration–drift equilibrium maintained by high gene flow. We found the highest rates of contemporary gene flow in populations that are connected by forest corridors. This information is highly relevant to conservation practitioners and policy makers, because deforestation, road widening and mining are imminent threats to these corridors.     

基于占域模型的中国珲春地区人虎冲突特征分析

人虎冲突是全球野生虎(Pantheratigris)保护和种群恢复工作迫切需要解决的问题,而人虎冲突的分布特征是开展冲突管理的重要科学依据.珲春地区是东北虎(P.t.altaica)从俄罗斯向中国扩散的重要廊道,随着东北虎种群数量的逐步增加,该地区的人虎冲突特征亟需开展研究.本文以2016-2020年珲春地区的人虎冲突...     

圈养华南虎的个性评估及其性别差异

个性研究在动物生态研究和迁地保护中发挥着重要作用.华南虎(Panthera tigris amoyensis)作为最濒危的虎亚种,目前仍然缺乏对其个性特征的研究探索.本文通过个性特征主观评价法对6只圈养华南虎进行个性评估,并对比不同性别的华南虎个性特征的差异,使用新异物体测验进一步验证个性特征主观评价法的结果.最终提取...     

Tigers of Sundarbans in India: Is the Population a Separate Conservation Unit?

The Sundarbans tiger inhabits a unique mangrove habitat and are morphologically distinct from the recognized tiger subspecies in terms of skull morphometrics and body size. Thus, there is an urgent need to assess their ecological and genetic distinctiveness and determine if Sundarbans tigers should be defined and managed as separate conservation unit. We utilized nine microsatellites and 3 kb from four mitochondrial DNA (mtDNA) genes to estimate genetic variability, population structure, demographic parameters and visualize historic and contemporary connectivity among tiger populations from Sundarbans and mainland India. We also evaluated the traits that determine exchangeability or adaptive differences among tiger populations. Data from both markers suggest that Sundarbans tiger is not a separate tiger subspecies and should be regarded as Bengal tiger (P. t. tigris) subspecies. Maximum likelihood phylogenetic analyses of the mtDNA data revealed reciprocal monophyly. Genetic differentiation was found stronger for mtDNA than nuclear DNA. Microsatellite markers indicated low genetic variation in Sundarbans tigers (He= 0.58) as compared to other mainland populations, such as northern and Peninsular (Hebetween 0.67- 0.70). Molecular data supports migration between mainland and Sundarbans populations until very recent times. We attribute this reduction in gene flow to accelerated fragmentation and habitat alteration in the landscape over the past few centuries. Demographic analyses suggest that Sundarbans tigers have diverged recently from peninsular tiger population within last 2000 years. Sundarbans tigers are the most divergent group of Bengal tigers, and ecologically non-exchangeable with other tiger populations, and thus should be managed as a separate “evolutionarily significant unit” (ESU) following the adaptive evolutionary conservation (AEC) concept.     

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.     

吉林珲春自然保护区东北虎捕食家畜研究

东北虎在中国已经极度濒危,随着位于中俄朝三国边境地区的吉林珲春自然保护区在2001 年底的建成,对虎的保护初见成效。然而保护区内虎频繁捕食家畜,引发了当地居民的不满。为了减缓人虎冲突,寻找东北虎捕食家畜的有关规律,作者开展了东北虎捕食家畜的相关研究。从2001 年12 月至2007 年底,共计有126 起,204 头家养动物被虎袭击,且呈现逐年上升的趋势。家畜被袭击主要发生在每年的4 ～ 9 月,没有发现明显的对家畜种的偏好。利用GIS 和回归分析表明:1)虎通常不袭击离开村庄小于1 km 的家畜;2)人为干扰对家畜的被食用率和虎移动家畜尸体的行为都有显著影响;3)共计有约16 079 kg 的家养动物肉被虎食用,造成损失76084 美元;4)对虎袭击地点生态变量的分析表明,东向和南向有更多的家畜被袭击;5)此外,虎多在较低的海拔和坡度、较高的郁闭度,靠近村庄和河流,远离道路等区域捕食家畜。清除套子和防止下套以增加有蹄类的密度对于减少虎捕食家畜非常重要。此外,减少虎袭击地点的人为干扰和改善对家畜的看护也非常关键。     

Prioritizing Tiger Conservation through Landscape Genetics and Habitat Linkages

Even with global support for tiger (Panthera tigris) conservation their survival is threatened by poaching, habitat loss and isolation. Currently about 3,000 wild tigers persist in small fragmented populations within seven percent of their historic range. Identifying and securing habitat linkages that connect source populations for maintaining landscape-level gene flow is an important long-term conservation strategy for endangered carnivores. However, habitat corridors that link regional tiger populations are often lost to development projects due to lack of objective evidence on their importance. Here, we use individual based genetic analysis in combination with landscape permeability models to identify and prioritize movement corridors across seven tiger populations within the Central Indian Landscape. By using a panel of 11 microsatellites we identified 169 individual tigers from 587 scat and 17 tissue samples. We detected four genetic clusters within Central India with limited gene flow among three of them. Bayesian and likelihood analyses identified 17 tigers as having recent immigrant ancestry. Spatially explicit tiger occupancy obtained from extensive landscape-scale surveys across 76,913 km² of forest habitat was found to be only 21,290 km². After accounting for detection bias, the covariates that best explained tiger occupancy were large, remote, dense forest patches; large ungulate abundance, and low human footprint. We used tiger occupancy probability to parameterize habitat permeability for modeling habitat linkages using least-cost and circuit theory pathway analyses. Pairwise genetic differences (F _ST) between populations were better explained by modeled linkage costs (r>0.5, p<0.05) compared to Euclidean distances, which was in consonance with observed habitat fragmentation. The results of our study highlight that many corridors may still be functional as there is evidence of contemporary migration. Conservation efforts should provide legal status to corridors, use smart green infrastructure to mitigate development impacts, and restore habitats where connectivity has been lost.     

Glucocorticoid Stress Responses of Reintroduced Tigers in Relation to Anthropogenic Disturbance in Sariska Tiger Reserve in India

Tiger (Panthera tigris), an endangered species, is under severe threat from poaching, habitat loss, prey depletion and habitat disturbance. Such factors have been reported causing local extermination of tiger populations including in one of the most important reserves in India, namely Sariska Tiger Reserve (STR) in northwestern India. Consequently, tigers were reintroduced in STR between 2008 and 2010, but inadequate breeding success was observed over the years, thus invoking an investigation to ascertain physiological correlates. In the present study, we report glucocorticoid stress responses of the reintroduced tigers in relation to anthropogenic disturbance in the STR from 2011 to 2013. We found anthropogenic disturbance such as encounter rates of livestock and humans, distance to roads and efforts to kill domestic livestock associated with an elevation in fecal glucocorticoid metabolite (fGCM) concentrations in the monitored tigers. In this regard, female tigers seem more sensitive to such disturbance than males. It was possible to discern that tiger’s fGCM levels were significantly positively related to the time spent in disturbed areas. Resulting management recommendations include relocation of villages from core areas and restriction of all anthropogenic activities in the entire STR.     

MHC class I and MHC class II DRB gene variability in wild and captive Bengal tigers (Panthera tigris tigris)

Bengal tigers are highly endangered and knowledge on adaptive genetic variation can be essential for efficient conservation and management. Here we present the first assessment of allelic variation in major histocompatibility complex (MHC) class I and MHC class II DRB genes for wild and captive tigers from India. We amplified, cloned, and sequenced alpha-1 and alpha-2 domain of MHC class I and beta-1 domain of MHC class II DRB genes in 16 tiger specimens of different geographic origin. We detected high variability in peptide-binding sites, presumably resulting from positive selection. Tigers exhibit a low number of MHC DRB alleles, similar to other endangered big cats. Our initial assessment—admittedly with limited geographic coverage and sample size—did not reveal significant differences between captive and wild tigers with regard to MHC variability. In addition, we successfully amplified MHC DRB alleles from scat samples. Our characterization of tiger MHC alleles forms a basis for further in-depth analyses of MHC variability in this illustrative threatened mammal.     

Why the Indian Subcontinent Holds the Key to Global Tiger Recovery

With only ~3,000 wild individuals surviving restricted to just 7% of their historical range, tigers are now a globally threatened species. Therefore, conservation efforts must prioritize regions that harbor more tigers, as well try to capture most of the remaining genetic variation and habitat diversity. Only such prioritization based on demographic, genetic, and ecological considerations can ensure species recovery and retention of evolutionary flexibility in the face of ongoing global changes. Although scientific understanding of ecological and demographic aspects of extant wild tiger populations has improved recently, little is known about their genetic composition and variability. We sampled 73 individual tigers from 28 reserves spread across a diversity of habitats in the Indian subcontinent to obtain 1,263 bp of mitochondrial DNA and 10 microsatellite loci. Our analyses reveals that Indian tigers retain more than half of the extant genetic diversity in the species. Coalescent simulations attribute this high genetic diversity to a historically large population size of about 58,200 tigers for peninsular India south of the Gangetic plains. Furthermore, our analyses indicate a precipitous, possibly human-induced population crash ~200 years ago in India, which is in concordance with historical records. Our results suggest that only 1.7% (with an upper limit of 13% and a lower limit of 0.2%) of tiger numbers in historical times remain now. In the global conservation context our results suggest that, based on genetic, demographic, and ecological considerations, the Indian subcontinent holds the key to global survival and recovery of wild tigers.     

Mitochondrial Phylogeography Illuminates the Origin of the Extinct Caspian Tiger and Its Relationship to the Amur Tiger

The Caspian tiger (Panthera tigris virgata) flourished in Central Asian riverine forest systems in a range disjunct from that of other tigers, but was driven to extinction in 1970 prior to a modern molecular evaluation. For over a century naturalists puzzled over the taxonomic validity, placement, and biogeographic origin of this enigmatic animal. Using ancient-DNA (aDNA) methodology, we generated composite mtDNA haplotypes from twenty wild Caspian tigers from throughout their historic range sampled from museum collections. We found that Caspian tigers carry a major mtDNA haplotype differing by only a single nucleotide from the monomorphic haplotype found across all contemporary Amur tigers (P. t. altaica). Phylogeographic analysis with extant tiger subspecies suggests that less than 10,000 years ago the Caspian/Amur tiger ancestor colonized Central Asia via the Gansu Corridor (Silk Road) from eastern China then subsequently traversed Siberia eastward to establish the Amur tiger in the Russian Far East. The conservation implications of these findings are far reaching, as the observed genetic depletion characteristic of modern Amur tigers likely reflects these founder migrations and therefore predates human influence. Also, due to their evolutionary propinquity, living Amur tigers offer an appropriate genetic source should reintroductions to the former range of the Caspian tiger be implemented.     

Genotyping faecal samples of Bengal tiger <Emphasis Type="Italic">Panthera tigris tigris</Emphasis> for population estimation: A pilot study

Background

Bengal tiger Panthera tigris tigris the National Animal of India, is an endangered species. Estimating populations for such species is the main objective for designing conservation measures and for evaluating those that are already in place. Due to the tiger's cryptic and secretive behaviour, it is not possible to enumerate and monitor its populations through direct observations; instead indirect methods have always been used for studying tigers in the wild. DNA methods based on non-invasive sampling have not been attempted so far for tiger population studies in India. We describe here a pilot study using DNA extracted from faecal samples of tigers for the purpose of population estimation.

Results

In this study, PCR primers were developed based on tiger-specific variations in the mitochondrial cytochrome b for reliably identifying tiger faecal samples from those of sympatric carnivores. Microsatellite markers were developed for the identification of individual tigers with a sibling Probability of Identity of 0.005 that can distinguish even closely related individuals with 99.9% certainty. The effectiveness of using field-collected tiger faecal samples for DNA analysis was evaluated by sampling, identification and subsequently genotyping samples from two protected areas in southern India.

Conclusion

Our results demonstrate the feasibility of using tiger faecal matter as a potential source of DNA for population estimation of tigers in protected areas in India in addition to the methods currently in use.

     

Influence of connectivity, wild prey and disturbance on occupancy of tigers in the human-dominated western Terai Arc Landscape

Occupying only 7% of their historical range and confined to forested habitats interspersed in a matrix of human dominated landscapes, tigers (Panthera tigris) typify the problems faced by most large carnivores worldwide. With heads of governments of tiger range countries pledging to reverse the extinction process and setting a goal of doubling wild tiger numbers by 2022, achieving this target would require identifying existing breeding cores, potential breeding habitats and opportunities for dispersal. The Terai Arc Landscape (TAL) represents one region which has recently witnessed recovery of tiger populations following conservation efforts. In this study, we develop a spatially explicit tiger occupancy model with survey data from 2009–10 based on a priori knowledge of tiger biology and specific issues plaguing the western TAL (6,979 km²), which occurs in two disjunct units (Tiger Habitat Blocks; THBs). Although the overall occupancy of tigers was 0.588 (SE 0.071), our results clearly indicate that loss in functionality of a regional corridor has resulted in tigers now occupying 17.58% of the available habitat in THB I in comparison to 88.5% in THB II. The current patterns of occupancy were best explained by models incorporating the interactive effect of habitat blocks (AIC w = 0.883) on wild prey availability (AIC w = 0.742) and anthropogenic disturbances (AIC w = 0.143). Our analysis has helped identify areas of high tiger occupancy both within and outside existing protected areas, which highlights the need for a unified control of the landscape under a single conservation unit with the primary focus of managing tigers and associated wildlife. Finally, in the light of global conservation targets and recent legislations in India, our study assumes significance as we identify opportunities to secure (e.g. THB II) and increase (e.g. THB I) tiger populations in the landscape.     

Estimating the Potential Impact of Canine Distemper Virus on the Amur Tiger Population (Panthera tigris altaica) in Russia

Lethal infections with canine distemper virus (CDV) have recently been diagnosed in Amur tigers (Panthera tigris altaica), but long-term implications for the population are unknown. This study evaluates the potential impact of CDV on a key tiger population in Sikhote-Alin Biosphere Zapovednik (SABZ), and assesses how CDV might influence the extinction potential of other tiger populations of varying sizes. An individual-based stochastic, SIRD (susceptible-infected-recovered/dead) model was used to simulate infection through predation of infected domestic dogs, and/or wild carnivores, and direct tiger-to-tiger transmission. CDV prevalence and effective contact based on published and observed data was used to define plausible low- and high-risk infection scenarios. CDV infection increased the 50-year extinction probability of tigers in SABZ by 6.3% to 55.8% compared to a control population, depending on risk scenario. The most significant factors influencing model outcome were virus prevalence in the reservoir population(s) and its effective contact rate with tigers. Adjustment of the mortality rate had a proportional impact, while inclusion of epizootic infection waves had negligible additional impact. Small populations were found to be disproportionately vulnerable to extinction through CDV infection. The 50-year extinction risk in populations consisting of 25 individuals was 1.65 times greater when CDV was present than that of control populations. The effects of density dependence do not protect an endangered population from the impacts of a multi-host pathogen, such as CDV, where they coexist with an abundant reservoir presenting a persistent threat. Awareness of CDV is a critical component of a successful tiger conservation management policy.     

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。     

Population status of a cryptic top predator: an island-wide assessment of tigers in Sumatran rainforests

Large carnivores living in tropical rainforests are under immense pressure from the rapid conversion of their habitat. In response, millions of dollars are spent on conserving these species. However, the cost-effectiveness of such investments is poorly understood and this is largely because the requisite population estimates are difficult to achieve at appropriate spatial scales for these secretive species. Here, we apply a robust detection/non-detection sampling technique to produce the first reliable population metric (occupancy) for a critically endangered large carnivore; the Sumatran tiger (Panthera tigris sumatrae). From 2007–2009, seven landscapes were surveyed through 13,511 km of transects in 394 grid cells (17×17 km). Tiger sign was detected in 206 cells, producing a naive estimate of 0.52. However, after controlling for an unequal detection probability (where p = 0.13±0.017; ±S.E.), the estimated tiger occupancy was 0.72±0.048. Whilst the Sumatra-wide survey results gives cause for optimism, a significant negative correlation between occupancy and recent deforestation was found. For example, the Northern Riau landscape had an average deforestation rate of 9.8%/yr and by far the lowest occupancy (0.33±0.055). Our results highlight the key tiger areas in need of protection and have led to one area (Leuser-Ulu Masen) being upgraded as a ‘global priority’ for wild tiger conservation. However, Sumatra has one of the highest global deforestation rates and the two largest tiger landscapes identified in this study will become highly fragmented if their respective proposed roads networks are approved. Thus, it is vital that the Indonesian government tackles these threats, e.g. through improved land-use planning, if it is to succeed in meeting its ambitious National Tiger Recovery Plan targets of doubling the number of Sumatran tigers by 2022.