Analyzing tiger interaction and home range shifts using a time-geographic approach.

BACKGROUND: Interaction through movement can be used as a marker to understand and model interspecific and intraspecific species dynamics, and the collective behavior of animals sharing the same space. This research leverages the time-geography framework, commonly used in human movement research, to explore the dynamic patterns of interaction between Indochinese tigers (Panthera tigris corbeti) in the western forest complex (WEFCOM) in Thailand. METHODS: We propose and assess ORTEGA, a time-geographic interaction analysis method, to trace spatio-temporal interactions patterns and home range shifts among tigers. Using unique GPS tracking data of tigers in WEFCOM collected over multiple years, concurrent and delayed interaction patterns of tigers are investigated. The outcomes are compared for intraspecific tiger interaction across different genders, relationships, and life stages. Additionally, the performance of ORTEGA is compared to a commonly used proximity-based approach. RESULTS: Among the 67 tracked tigers, 42 show concurrent interactions at shared boundaries. Further investigation of five tigers with overlapping home ranges (two adult females, a male, and two young male tigers) suggests that the mother tiger and her two young mostly stay together before their dispersal but interact less post-dispersal. The male tiger increases encounters with the mother tiger while her young shift their home ranges. On another timeline, the neighbor female tiger mostly avoids the mother tiger. Through these home range dynamics and interaction patterns, we identify four types of interaction among these tigers: following, encounter, latency, and avoidance. Compared to the proximity-based approach, ORTEGA demonstrates better detects concurrent mother-young interactions during pre-dispersal, while the proximity-based approach misses many interactions among the dyads. With larger spatial buffers and temporal windows, the proximity-based approach detects more encounters but may overestimate the duration of interaction. CONCLUSIONS: This research demonstrates the applicability and merits of ORTEGA as a time-geographic based approach to animal movement interaction analysis. We show time geography can develop valuable, data-driven insights about animal behavior and interactions. ORTEGA effectively traces frequent encounters and temporally delayed interactions between animals, without relying on specific spatial and temporal buffers. Future research should integrate contextual and behavioral information to better identify and characterize the nature of species interaction.

Competitive interactions and coexistence of sympatric flagship carnivores in Asia.

Understanding the competition and coexistence of flagship carnivores is key to creating strategies for their conservation in the face of global carnivore declines. Although studies exploring the dynamics and competition between tigers (Panthera tigris) and leopards (P. pardus) span decades, there is a lack of understanding regarding the factors that influence their coexistence mechanisms on a broad scale, as well as the drivers determining their exploitative and interference competition. We gathered a comprehensive list of research papers among which 36 papers explored the interspecific interactions between tigers and leopards and tested the influence of biotic and abiotic factors on the coexistence mechanisms along three dimensions using multiple response variables regression models; we also tested the influence of ecological drivers determining the exploitative or interference competition between tigers and leopards. Elevation and ungulate density were the most important predictors in regulating the coexistence mechanisms. Tigers and leopards exhibited more positive relations/higher overlaps as elevation increased in the spatial niche. In addition, they showed a higher dietary overlap in the prey-rich regions. We determined that interference competition between tigers and leopards was less frequently observed in habitats with dense tree cover and homogeneous vegetation structures. Meanwhile, studies with multiple metrics would promote the detection of interference competition. Our study provides new insight into the competitive interactions and coexistence mechanisms of tigers and leopards on a broad scale. Policy-makers and managers should pay more attention to the factors of elevation, prey abundance, and habitat structures for the conservation of tigers and leopards.

Addressing the impact of canine distemper spreading on an isolated tiger population in northeast Asia.

The continuation of the isolated Amur tiger (Panthera tigris altaica) population living along the China-Russia border is facing serious challenges due to factors such as its small size (including 38 individuals) and canine distemper virus (CDV). We use a population viability analysis metamodel, which consists of a traditional individual-based demographic model linked to an epidemiological model, to assess options for controlling the impact of negative factors through domestic dog management in protected areas, increasing connectivity to the neighboring large population (including more than 400 individuals), and habitat expansion. Without intervention, under inbreeding depression of 3.14, 6.29, and 12.26 lethal equivalents, our metamodel predicted the extinction within 100 years is 64.4%, 90.6%, and 99.8%, respectively. In addition, the simulation results showed that dog management or habitat expansion independently will not ensure tiger population viability for the next 100 years, and connectivity to the neighboring population would only keep the population size from rapidly declining. However, when the above three conservation scenarios are combined, even at the highest level of 12.26 lethal equivalents inbreeding depression, population size will not decline and the probability of extinction will be <5.8%. Our findings highlight that protecting the Amur tiger necessitates a multifaceted synergistic effort. Our key management recommendations for this population underline the importance of reducing CDV threats and expanding tiger occupancy to its former range in China, but re-establishing habitat connectivity to the neighboring population is an important long-term objective.

Contributions of distemper control and habitat expansion to the Amur leopard viability.

The Amur leopard (Panthera pardus orientalis) is a critically endangered top predator that struggles on the brink of extinction due to threats such as canine distemper virus (CDV), habitat loss, and inbreeding depression. Here we develop a viability analysis metamodel that combines a traditional individual-based demographic model with an epidemiological model to assess the benefits of alternative population management actions in response to multiple distinct threats. Our results showed an extinction risk of 10.3%-99.9% if no management actions were taken over 100 years under different levels of inbreeding depression. Reducing the risk of CDV infection in Amur leopards through the low-coverage vaccination of leopards and the management of sympatric domestic dogs could effectively improve the survival probability of the leopard population, and with habitat expansion added to these management measures, the population expanded further. Our findings highlight that protecting the Amur leopard necessitates a multifaceted synergistic effort, and controlling multiple threats together may significantly escalate overall viability of a species, especially for small-isolated threatened population. More broadly, our modeling framework could offer critical perspectives and scientific support for conservation planning, as well as specific adaptive management actions for endangered species around the world.

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.

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

Despite conservation efforts, large mammals such as tigers (Panthera tigris) and their main prey, gaur (Bos gaurus), banteng (Bos javanicus), and sambar (Rusa unicolor), are highly threatened and declining across their entire range. The only large viable source population of tigers in mainland Southeast Asia occurs in Thailand's Western Forest Complex (WEFCOM), an approximately 19,000 km2 landscape of 17 contiguous protected areas.We used an occupancy modeling framework, which accounts for imperfect detection, to identify the factors that affect tiger distribution at the approximate scale of a female tiger's home range, 64 km2, and site use at a scale of 1-km2. At the larger scale, we estimated the proportion of sites at WEFCOM that were occupied by tigers; at the finer scale, we identified the key variables that influence site-use and developed a predictive distribution map. At both scales, we examined key anthropogenic and ecological factors that help explain tiger distribution and habitat use, including probabilities of gaur, banteng, and sambar occurrence from a companion study.Occupancy estimated at the 64-km2 scale was primarily influenced by the combined presence of all three large prey species, and 37% or 5,858 km2 of the landscape was predicted to be occupied by tigers. In contrast, site use estimated at the scale of 1 km2 was most strongly influenced by the presence of sambar.By modeling occupancy while accounting for imperfect probability of detection, we established reliable benchmark data on the distribution of tigers in WEFCOM. This study also identified factors that limit tiger distributions; which managers can then target to expand tiger distribution and guide recovery elsewhere in Southeast Asia.

Genome-Wide Evolutionary Analysis of Natural History and Adaptation in the World's Tigers.

No other species attracts more international resources, public attention, and protracted controversies over its intraspecific taxonomy than the tiger (Panthera tigris) [1, 2]. Today, fewer than 4,000 free-ranging tigers survive, covering only 7% of their historical range, and debates persist over whether they comprise six, five, or two subspecies [3-6]. The lack of consensus over the number of tiger subspecies has partially hindered the global effort to recover the species from the brink of extinction, as both captive breeding and landscape intervention of wild populations increasingly require an explicit delineation of the conservation management units [7]. The recent coalescence to a late Pleistocene bottleneck (circa 110 kya) [5, 8, 9] poses challenges for detecting tiger subspecific morphological traits, suggesting that elucidating intraspecific evolution in the tiger requires analyses at the genomic scale. Here, we present whole-genome sequencing analyses from 32 voucher specimens that resolve six statistically robust monophyletic clades corresponding to extant subspecies, including the recently recognized Malayan tiger (P. tigris jacksoni). The intersubspecies gene flow is very low, corroborating the recognized phylogeographic units. We identified multiple genomic regions that are candidates for identifying the adaptive divergence of subspecies. The body-size-related gene ADH7 appears to have been strongly selected in the Sumatran tiger, perhaps in association with adaptation to the tropical Sunda Islands. The identified genomic signatures provide a solid basis for recognizing appropriate conservation management units in the tiger and can benefit global conservation strategic planning for this charismatic megafauna icon.

How much gene flow is needed to avoid inbreeding depression in wild tiger populations?

The number and size of tiger populations continue to decline owing to habitat loss, habitat fragmentation and poaching of tigers and their prey. As a result, tiger populations have become small and highly structured. Current populations have been isolated since the early 1970s or for approximately seven generations. The objective of this study is to explore how inbreeding may be affecting the persistence of remaining tiger populations and how dispersal, either natural or artificial, may reduce the potentially detrimental effect of inbreeding depression. We developed a tiger simulation model and used published levels of genetic load in mammals to simulate inbreeding depression. Following a 50 year period of population isolation, we introduced one to four dispersing male tigers per generation to explore how gene flow from nearby populations may reduce the negative impact of inbreeding depression. For the smallest populations, even four dispersing male tigers per generation did not increase population viability, and the likelihood of extinction is more than 90% within 30 years. Unless habitat connectivity is restored or animals are artificially introduced in the next 70 years, medium size wild populations are also likely to go extinct, with only four to five of the largest wild tiger populations likely to remain extant in this same period without intervention. To reduce the risk of local extinction, habitat connectivity must be pursued concurrently with efforts to increase population size (e.g. enhance habitat quality, increase habitat availability). It is critical that infrastructure development, dam construction and other similar projects are planned appropriately so that they do not erode the extent or quality of habitat for these populations so that they can truly serve as future source populations.

Sympatric Asian felid phylogeography reveals a major Indochinese-Sundaic divergence.

The dynamic geological and climatological history of Southeast Asia has spawned a complex array of ecosystems and 12 of the 37 known cat species, making it the most felid-rich region in the world. To examine the evolutionary histories of these poorly studied fauna, we compared phylogeography of six species (leopard cat Prionailurus bengalensis, fishing cat P. viverrinus, Asiatic golden cat Pardofelis temminckii, marbled cat P. marmorata, tiger Panthera tigris and leopard P. pardus) by sequencing over 5 kb of DNA each from 445 specimens at multiple loci of mtDNA, Y and X chromosomes. All species except the leopard displayed significant phylogenetic partitions between Indochina and Sundaland, with the central Thai-Malay Peninsula serving as the biogeographic boundary. Concordant mtDNA and nuclear DNA genealogies revealed deep Indochinese-Sundaic divergences around 2 MYA in both P. bengalensis and P. marmorata comparable to previously described interspecific distances within Felidae. The divergence coincided with serial sea level rises during the late Pliocene and early Pleistocene, and was probably reinforced by repeated isolation events associated with environmental changes throughout the Pleistocene. Indochinese-Sundaic differentiations within P. tigris and P. temminckii were more recent at 72-108 and 250-1570 kya, respectively. Overall, these results illuminate unexpected, deep vicariance events in Southeast Asian felids and provide compelling evidence of species-level distinction between the Indochinese and Sundaic populations in the leopard cat and marbled cat. Broader sampling and further molecular and morphometric analyses of these species will be instrumental in defining conservation units and effectively preserving Southeast Asian biodiversity.

Use of an action-selection framework for human-carnivore conflict in the Bangladesh Sundarbans.

Human-carnivore conflict is manifested in the death of humans, livestock, and carnivores. The resulting negative local attitudes and retribution killings imperil the future of many endangered carnivores. We tailored existing management tools to create a framework to facilitate the selection of actions to alleviate human-carnivore conflict and applied the framework to the human-tiger conflict in the Bangladesh Sundarbans. We identified potential actions that consider previous management efforts, local knowledge, cost-effectiveness, fieldwork experience of authors and project staff, previous research on tiger ecology by the authors, and recommendations from human-carnivore conflict studies in other countries. Our framework includes creation of a profile to improve understanding of the nature of the conflict and its underlying causality. Identified actions include deterrents, education, direct tiger management, and response teams. We ranked actions by their potential to reduce conflict and the monetary cost of their implementation. We ranked tiger-response teams and monitoring problem tigers as the two best actions because both had relatively high impact and cost-effectiveness. We believe this framework could be used under a wide range of human-wildlife conflict situations because it provides a structured approach to selection of mitigating actions.

Subspecies genetic assignments of worldwide captive tigers increase conservation value of captive populations.

Tigers (Panthera tigris) are disappearing rapidly from the wild, from over 100,000 in the 1900s to as few as 3000. Javan (P.t. sondaica), Bali (P.t. balica), and Caspian (P.t. virgata) subspecies are extinct, whereas the South China tiger (P.t. amoyensis) persists only in zoos. By contrast, captive tigers are flourishing, with 15,000-20,000 individuals worldwide, outnumbering their wild relatives five to seven times. We assessed subspecies genetic ancestry of 105 captive tigers from 14 countries and regions by using Bayesian analysis and diagnostic genetic markers defined by a prior analysis of 134 voucher tigers of significant genetic distinctiveness. We assigned 49 tigers to one of five subspecies (Bengal P.t. tigris, Sumatran P.t. sumatrae, Indochinese P.t. corbetti, Amur P.t. altaica, and Malayan P.t. jacksoni tigers) and determined 52 had admixed subspecies origins. The tested captive tigers retain appreciable genomic diversity unobserved in their wild counterparts, perhaps a consequence of large population size, century-long introduction of new founders, and managed-breeding strategies to retain genetic variability. Assessment of verified subspecies ancestry offers a powerful tool that, if applied to tigers of uncertain background, may considerably increase the number of purebred tigers suitable for conservation management.

Development of Y chromosome intraspecific polymorphic markers in the Felidae.

Y chromosome haplotyping based on microsatellites and single nucleotide polymorphisms (SNPs) has proved to be a powerful tool for population genetic studies of humans. However, the promise of the approach is hampered in the majority of nonhuman mammals by the lack of Y-specific polymorphic markers. We were able to identify new male-specific polymorphisms in the domestic cat Felis catus and 6 additional Felidae species with a combination of molecular genetic and cytogenetic approaches including 1) identifying domestic cat male-specific microsatellites from markers generated from a male cat microsatellite-enriched genomic library, a flow-sorted Y cosmid library, or a Y-specific cat bacteria artificial chromosome (BAC) clone, (2) constructing microsatellite-enriched libraries from flow-sorted Y chromosomes isolated directly from focal wildcat species, and (3) screening Y chromosome conserved anchored tagged sequences primers in Felidae species. Forty-one male-specific microsatellites were identified, but only 6 were single-copy loci, consistent with the repetitive nature of the Y chromosome. Nucleotide diversity (pi) of Y-linked intron sequences (2.1 kbp) was in the range of 0 (tiger) to 9.95 x 10(-4) (marbled cat), and the number of SNPs ranged from none in the tiger to 7 in the Asian leopard cat. The Y haplotyping system described here, consisting of 4 introns (SMCY3, SMCY7, UTY11, and DBY7) and 1 polymorphic microsatellite (SMCY-STR), represents the first available markers for tracking intraspecific male lineage polymorphisms in Felidae species and promises to provide significant insights to evolutionary and population genetic studies of the species.

Phylogeography and genetic ancestry of tigers (Panthera tigris).

Eight traditional subspecies of tiger (Panthera tigris),of which three recently became extinct, are commonly recognized on the basis of geographic isolation and morphological characteristics. To investigate the species' evolutionary history and to establish objective methods for subspecies recognition, voucher specimens of blood, skin, hair, and/or skin biopsies from 134 tigers with verified geographic origins or heritage across the whole distribution range were examined for three molecular markers: (1) 4.0 kb of mitochondrial DNA (mtDNA) sequence; (2) allele variation in the nuclear major histocompatibility complex class II DRB gene; and (3) composite nuclear microsatellite genotypes based on 30 loci. Relatively low genetic variation with mtDNA,DRB,and microsatellite loci was found, but significant population subdivision was nonetheless apparent among five living subspecies. In addition, a distinct partition of the Indochinese subspecies P. t. corbetti in to northern Indochinese and Malayan Peninsula populations was discovered. Population genetic structure would suggest recognition of six taxonomic units or subspecies: (1) Amur tiger P. t. altaica; (2) northern Indochinese tiger P. t. corbetti; (3) South China tiger P. t. amoyensis; (4) Malayan tiger P. t. jacksoni, named for the tiger conservationist Peter Jackson; (5) Sumatran tiger P. t. sumatrae; and (6) Bengal tiger P. t. tigris. The proposed South China tiger lineage is tentative due to limited sampling. The age of the most recent common ancestor for tiger mtDNA was estimated to be 72,000-108,000 y, relatively younger than some other Panthera species. A combination of population expansions, reduced gene flow, and genetic drift following the last genetic diminution, and the recent anthropogenic range contraction, have led to the distinct genetic partitions. These results provide an explicit basis for subspecies recognition and will lead to the improved management and conservation of these recently isolated but distinct geographic populations of tigers.

The Long-Term Effects of Tiger Poaching on Population Viability.

Poaching tigers, primarily for their bones, has become the latest threat to the persistence of wild tiger populations throughout the world. Anecdotal information indicates the seriousness of this new threat. It is important, however, to provide a quantitative analysis of poaching as a basis for strong policy action. We therefore created a tiger simulation model to explore the effects of realistic levels of poaching on population viability. The model is an individually based, stochastic spatial model that is based on the extensive data set from Royal Chitwan National Park, Nepal. We found that as poaching continues over time, the probability of population extinction increases sigmoidally; a critical zone exists in which a small, incremental increase in poaching greatly increases the probability of extinction. The implication is that poaching may not at first be seen as a threat but could suddenly become one. Moreover, even if poaching is effectively stopped, tiger populations will still be vulnerable and could go extinct due to demographic and environmental stochasticity. Our model also shows that poaching reduces genetic variability, which could further reduce population viability due to inbreeding depression. The longer poaching is allowed to continue, the more vulnerable a population will be to these stochastic events. At currently reported rates of poaching our analysis indicates that many wild tiger populations will be extirpated during the latter half of the 1990s. Los efectos a largo plazo de la caza furtiva de tigres sobre la viabilidad poblacional.

Resumen: La caza furtiva de tigres, principalmente por sus huesos, se ha transformado en la última amenaza para la persistencia de problaciones salvajes de tigres alrededor del mundo. Información anecdótica indica la seriedad de esta nueva amenaza. Es importante, sin embargo proveer un análisis cuantitativo de la caza furtiva como base para una acción política firme. Es por ello que nosotros creamos un modelo de simulación para explorar los efectos de niveles realistas de caza furtiva sobre la viabilidad poblacional. Usamos un modelo espacial estocástico basado en individuos, que esta basado en el extenso conjunto de datos del Parque Nacional Real de Chitwan, en Nepal. Encontramos que a medida que la caza furtiva continúa, la probabilidad de extinción poblacional aumenta sigmoidalmente; existe una zona crítica donde un pequeño incremento en la caza furtiva incrementa en gran medida la probabilidad de extinción. La complicación es que la caza furtiva podría no ser vista al principio como una amenaza, pero pondría de pronto transformarse en una. Más aún, si la caza furtiva es detenida eficientemente, las poblaciones de tigres serían todavia vulnerables y podrían extinguirse debido a la estocasticidad demográfica y ambiental. Nuestro modelo también demuestra que la caza furtiva reduce la variabilidad genética, lo cual podria reducir aún más la viabilidad poblacional debido a la depresión de endocría. Cuanto más tiempo se permita que la caza furtiva continúe, más vulnerables se harán las poblaciones a estos eventos stocásticos. Nuestro análisis indica que al ritmo de las tasas de caza furtiva reportadas en la actualidad, muchas poblaciones de tigres serían eliminadas durante la última mitad de la década de los 90.