Distemper, extinction, and vaccination of the Amur tiger.

Canine distemper virus (CDV) has recently emerged as an extinction threat for the endangered Amur tiger (Panthera tigris altaica). CDV is vaccine-preventable, and control strategies could require vaccination of domestic dogs and/or wildlife populations. However, vaccination of endangered wildlife remains controversial, which has led to a focus on interventions in domestic dogs, often assumed to be the source of infection. Effective decision making requires an understanding of the true reservoir dynamics, which poses substantial challenges in remote areas with diverse host communities. We carried out serological, demographic, and phylogenetic studies of dog and wildlife populations in the Russian Far East to show that a number of wildlife species are more important than dogs, both in maintaining CDV and as sources of infection for tigers. Critically, therefore, because CDV circulates among multiple wildlife sources, dog vaccination alone would not be effective at protecting tigers. We show, however, that low-coverage vaccination of tigers themselves is feasible and would produce substantive reductions in extinction risks. Vaccination of endangered wildlife provides a valuable component of conservation strategies for endangered species.

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.

Assessing the health risks of reintroduction: The example of the Amur leopard, Panthera pardus orientalis.

Translocation of wildlife as a means of reintroducing or reinforcing threatened populations is an important conservation tool but carries health risks for the translocated animals and their progeny, as well as wildlife, domestic animals and humans in the release area. Disease risk analyses (DRA) are used to identify, prioritize and design mitigation strategies to address these threats. Here, we use a DRA undertaken for Amur leopards (Panthera pardus orientalis) to illustrate how specific methodology can optimize mitigation strategy design. A literature review identified a total of 98 infectious hazards and 28 non-infectious hazards. Separate analyses were undertaken for disease risks in leopards from hazards of source origin (captive zoo collections and the transit pathway to the Russian Far East), or of destination origin (in breeding enclosures and wider release areas); and for disease risks in other wildlife, domesticated species or humans, similarly from hazards of source or destination origin. Hazards were assessed and ranked as priority 1, priority 2, priority 3 or low priority in each of the defined scenarios. In addition, we undertook a generic assessment of stress on individual leopards. We use three examples to illustrate the process: Chlamydophila felis, canine distemper virus (CDV) and feline immunodeficiency virus (FIV). We found that many potentially expensive screening procedures could be performed prior to export of leopards, putting the onus of responsibility onto the zoo sector, for which access to diagnostic testing facilities is likely to be optimal. We discuss how our methods highlighted significant data gaps relating to pathogen prevalence in the Russian Far East and likely future unpredictability, in particular with respect to CDV. There was emphasis at all stages on record keeping, meticulous planning, design, staff training and enclosure management, which are relatively financially inexpensive. Actions to minimize stress featured at all time points in the strategy and also focussed on planning, design and management.

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.

CANINE DISTEMPER VIRUS IN A WILD FAR EASTERN LEOPARD ( PANTHERA PARDUS ORIENTALIS).

The critically endangered population of Far Eastern leopards ( Panthera pardus orientalis) may number as few as 60 individuals and is at risk from stochastic processes such as infectious disease. During May 2015, a case of canine distemper virus (CDV) was diagnosed in a wild leopard exhibiting severe neurologic disease in the Russian territory of Primorskii Krai. Amplified sequences of the CDV hemagglutinin gene and phosphoprotein gene aligned within the Arctic-like clade of CDV, which includes viruses from elsewhere in Russia, China, Europe, and North America. Histologic examination of cerebral tissue revealed perivascular lymphoid cuffing and demyelination of the white matter consistent with CDV infection. Neutralizing antibodies against CDV were detected in archived serum from two wild Far Eastern leopards sampled during 1993-94, confirming previous exposure in the population. This leopard population is likely too small to maintain circulation of CDV, suggesting that infections arise from spillover from more-abundant domestic or wild carnivore reservoirs. Increasing the population size and establishment of additional populations of leopards would be important steps toward securing the future of this subspecies and reducing the risk posed by future outbreaks of CDV or other infectious diseases.

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.

Mapping black panthers: Macroecological modeling of melanism in leopards (Panthera pardus).

The geographic distribution and habitat association of most mammalian polymorphic phenotypes are still poorly known, hampering assessments of their adaptive significance. Even in the case of the black panther, an iconic melanistic variant of the leopard (Panthera pardus), no map exists describing its distribution. We constructed a large database of verified records sampled across the species' range, and used it to map the geographic occurrence of melanism. We then estimated the potential distribution of melanistic and non-melanistic leopards using niche-modeling algorithms. The overall frequency of melanism was ca. 11%, with a significantly non-random spatial distribution. Distinct habitat types presented significantly different frequencies of melanism, which increased in Asian moist forests and approached zero across most open/dry biomes. Niche modeling indicated that the potential distributions of the two phenotypes were distinct, with significant differences in habitat suitability and rejection of niche equivalency between them. We conclude that melanism in leopards is strongly affected by natural selection, likely driven by efficacy of camouflage and/or thermoregulation in different habitats, along with an effect of moisture that goes beyond its influence on vegetation type. Our results support classical hypotheses of adaptive coloration in animals (e.g. Gloger's rule), and open up new avenues for in-depth evolutionary analyses of melanism in mammals.

Estimating abundance and density of Amur tigers along the Sino-Russian border.

As an apex predator the Amur tiger (Panthera tigris altaica) could play a pivotal role in maintaining the integrity of forest ecosystems in Northeast Asia. Due to habitat loss and harvest over the past century, tigers rapidly declined in China and are now restricted to the Russian Far East and bordering habitat in nearby China. To facilitate restoration of the tiger in its historical range, reliable estimates of population size are essential to assess effectiveness of conservation interventions. Here we used camera trap data collected in Hunchun National Nature Reserve from April to June 2013 and 2014 to estimate tiger density and abundance using both maximum likelihood and Bayesian spatially explicit capture-recapture (SECR) methods. A minimum of 8 individuals were detected in both sample periods and the documentation of marking behavior and reproduction suggests the presence of a resident population. Using Bayesian SECR modeling within the 11 400 km(2) state space, density estimates were 0.33 and 0.40 individuals/100 km(2) in 2013 and 2014, respectively, corresponding to an estimated abundance of 38 and 45 animals for this transboundary Sino-Russian population. In a maximum likelihood framework, we estimated densities of 0.30 and 0.24 individuals/100 km(2) corresponding to abundances of 34 and 27, in 2013 and 2014, respectively. These density estimates are comparable to other published estimates for resident Amur tiger populations in the Russian Far East. This study reveals promising signs of tiger recovery in Northeast China, and demonstrates the importance of connectivity between the Russian and Chinese populations for recovering tigers in Northeast China.

Detection of Hepatozoon felis in Ticks Collected from Free-Ranging Amur Tigers ( Panthera tigris altaica), Russian Far East, 2002-12.

We collected 69 ticks from nine, free-ranging Amur tigers ( Panthera tigris altaica) between 2002 and 2011 and investigated them for tick-borne pathogens. DNA was extracted using alkaline digestion and PCR was performed to detect apicomplexan organisms. Partial 18S rDNA amplification products were obtained from 14 ticks from four tigers, of which 13 yielded unambiguous nucleotide sequence data. Comparative sequence analysis revealed all 13 partial 18S rDNA sequences were most similar to those belonging to strains of Hepatozoon felis (>564/572 base-pair identity, >99% sequence similarity). Although this tick-borne protozoon pathogen has been detected in wild felids from many parts of the world, this is the first record from the Russian Far East.

Indicators of success for smart law enforcement in protected areas: A case study for Russian Amur tiger (Panthera tigris altaica) reserves.

Although considerable conservation resources have been committed to develop and use law enforcement monitoring and management tools such as SMART, measures of success are ill-defined and, to date, few reports detail results post-implementation. Here, we present 4 case studies from protected areas with Amur tigers (Panthera tigris altaica) in Russia, in which indicators of success were defined and evaluated at each. The ultimate goal was an increase in tiger numbers to 1 individual/100 km(2) at each site. We predicted that improvements in law enforcement effectiveness would be followed by increases in prey numbers and, subsequently, tiger numbers. We used short-term and long-term indicators of success, including: (i) patrol team effort and effectiveness; (ii) catch per unit effort indicators (to measure reductions in threats); and (iii) changes in target species numbers. In addition to implementing a monitoring system, we focused on improving law enforcement management using an adaptive management process. Over 4 years, we noted clear increases in patrol effort and a partial reduction in threats. Although we did not detect clear trends in ungulate numbers, tiger populations remained stable or increased, suggesting that poaching of tigers may be more limiting than prey depletion. Increased effectiveness is needed before a clear reduction in threats can be noted, and more time is needed before detecting responses in target populations. Nonetheless, delineation of concrete goals and indicators of success provide a means of evaluating progress and weaknesses. Such monitoring should be a central component of law enforcement strategies for protected areas.

Comparison of carnivore, omnivore, and herbivore mammalian genomes with a new leopard assembly.

BACKGROUND: There are three main dietary groups in mammals: carnivores, omnivores, and herbivores. Currently, there is limited comparative genomics insight into the evolution of dietary specializations in mammals. Due to recent advances in sequencing technologies, we were able to perform in-depth whole genome analyses of representatives of these three dietary groups. RESULTS: We investigated the evolution of carnivory by comparing 18 representative genomes from across Mammalia with carnivorous, omnivorous, and herbivorous dietary specializations, focusing on Felidae (domestic cat, tiger, lion, cheetah, and leopard), Hominidae, and Bovidae genomes. We generated a new high-quality leopard genome assembly, as well as two wild Amur leopard whole genomes. In addition to a clear contraction in gene families for starch and sucrose metabolism, the carnivore genomes showed evidence of shared evolutionary adaptations in genes associated with diet, muscle strength, agility, and other traits responsible for successful hunting and meat consumption. Additionally, an analysis of highly conserved regions at the family level revealed molecular signatures of dietary adaptation in each of Felidae, Hominidae, and Bovidae. However, unlike carnivores, omnivores and herbivores showed fewer shared adaptive signatures, indicating that carnivores are under strong selective pressure related to diet. Finally, felids showed recent reductions in genetic diversity associated with decreased population sizes, which may be due to the inflexible nature of their strict diet, highlighting their vulnerability and critical conservation status. CONCLUSIONS: Our study provides a large-scale family level comparative genomic analysis to address genomic changes associated with dietary specialization. Our genomic analyses also provide useful resources for diet-related genetic and health research.

Population dynamics of Amur tigers (Panthera tigris altaica) in Sikhote-Alin Biosphere Zapovednik: 1966-2012.

In 2010, the world's tiger (Panthera tigris) range countries agreed to the goal of doubling tiger numbers over 12 years, but whether such an increase is biologically feasible has not been assessed. Long-term monitoring of tigers in Sikhote-Alin Biosphere Zapovednik (SABZ), Russia provided an opportunity to determine growth rates of a recovering population. A 41-year growth phase was followed by a rapid decline in tiger numbers. Annual growth rates during the growth phase averaged 4.6%, beginning near 10% in the earliest years but quickly dropping below 5%. Sex ratio (females per male) mirrored growth rates, declining as population size increased. The rapid decline from 2009 to 2012 appeared to be tied to multiple factors, including poaching, severe winters and disease. Reproductive indicators of this population are similar to those of Bengal tiger populations, suggesting that growth rates may be similar. These results suggest that, first, tiger populations likely in general grow slowly: 3-5% yearly increases are realistic and larger growth rates are likely only when populations are highly depressed, mortality rates are low and prey populations are high relative to numbers of adult females. Second, while more research is needed, it should not be assumed that tiger populations with high prey densities will necessarily grow more quickly than populations with low prey densities. Third, while growth is slow, decline can be rapid. Fourth, because declines can happen so quickly, there is a constant need to monitor populations and be ready to respond with appropriate and timely conservation interventions if tiger populations are to remain secure. Finally, an average annual growth rate across all tiger populations of 6%, required to reach the Global Tiger Initiative's goal of doubling tiger numbers in 12 years, is a noble but unlikely scenario.

Mortality of Amur tigers: The more things change, the more they stay the same.

Poaching as well as loss of habitat and prey are identified as causes of tiger population declines. Although some studies have examined habitat requirements and prey availability, few studies have quantified cause-specific mortality of tigers. We used cumulative incidence functions (CIFs) to quantify cause-specific mortality rates of tigers, expanding and refining earlier studies to assess the potential impact of a newly emerging disease. To quantify changes in tiger mortality over time, we re-examined data first collected by Goodrich et al. (; study period 1: 1992-2004) as well as new telemetry data collected since January 2005 (study period 2: 2005-2012) using a total of 57 tigers (27 males and 30 females) monitored for an average of 747 days (range 26-4718 days). Across the entire study period (1992 to 2012) we found an estimated average annual survival rate of 0.75 for all tigers combined. Poaching was the primary cause of mortality during both study periods, followed by suspected poaching, distemper and natural/unknown causes. Since 2005, poaching mortality has remained relatively constant and, if combined with suspected poaching, may account for a loss of 17-19% of the population each year. Canine distemper virus (CDV) may be an additive form of mortality to the population, currently accounting for an additional 5%. Despite this relatively new source of mortality, poaching remains the main threat to Amur tiger survival and, therefore, population growth.

Identifying ecological corridors for Amur tigers (Panthera tigris altaica) and Amur leopards (Panthera pardus orientalis).

The rapid explosion of human populations and the associated development of human-dominated landscapes have drastically reduced and fragmented habitat for tigers (Panthera tigris) and leopards (Panthera pardus) across Asia, resulting in multiple small populations. However, Amur tiger (Panthera tigris altaica) habitat in Russia has remained largely interconnected, except for a break between tigers in southwest Primorye and the southern Sikhote-Alin Mountains. This habitat patch in southwest Primorye also retains the last population of Amur leopards (Panthera pardus orientalis). Genetic differentiation of tigers in southwest Primorye and the Sikhote-Alin Mountains along with survey data suggest that habitat fragmentation is limiting movement of tigers and leopards across the Razdolnaya River basin. We looked at historical and recent survey data on tigers and leopards and mapped existing cover types to examine land-use patterns of both large felids and humans in the development strip along the Razdolnaya River. We then used least-cost distance analyses to identify the most effective potential corridor to retain connectivity for large felids between Land of the Leopard National Park and Ussuriskii Zapovednik (Reserve). We identified a single potential corridor that still exists with a total distance of 62.5 km from Land of the Leopard National Park to Ussuriskii Zapovednik, mostly (93%) through forested habitat. We recommend formal recognition of a Razdolnaya ecological corridor and provide specific recommendations for each of 3 proposed management sections.

A comparison of food habits and prey preference of Amur tiger (Panthera tigris altaica) at three sites in the Russian Far East.

Prey availability is one of the principal drivers of tiger distribution and abundance. Therefore, formulating effective conservation strategies requires a clear understanding of tiger diet. We used scat analysis in combination with data on the abundance of several prey species to estimate Amur tiger diet and preference at 3 sites in the Russian Far East. We also examined the effect of pseudoreplication on estimates of tiger diet. We collected 770 scats across the 3 sites. Similar to previous studies, we found that tigers primarily preyed on medium to large ungulates, with wild boar, roe, sika and red deer collectively comprising 86.7% of total biomass consumed on average. According to Jacobs' index, tigers preferred wild boar, and avoided sika deer. Variation in preference indices derived from these scat analyses compared to indices derived from kill data appear to be due to adjustments in biomass intake when sex-age of a killed individual is known: a component missing from scat data. Pseudoreplication (multiple samples collected from a single kill site) also skewed results derived from scat analyses. Scat analysis still appears useful in providing insight into the diets of carnivores when the full spectrum of prey species needs to be identified, or when sample sizes from kill data are not sufficient. When sample sizes of kill data are large (as is now possible with GPS-collared animals), kill data adjusted by sex-age categories probably provides the most accurate estimates of prey biomass composition. Our results provide further confirmation of the centrality of medium ungulates, in particular wild boar, to Amur tiger diet, and suggest that the protection of this group of species is critical to Amur tiger conservation.

Canine distemper virus as a threat to wild tigers in Russia and across their range.

Canine distemper virus (CDV) has recently been identified in populations of wild tigers in Russia and India. Tiger populations are generally too small to maintain CDV for long periods, but are at risk of infections arising from more abundant susceptible hosts that constitute a reservoir of infection. Because CDV is an additive mortality factor, it could represent a significant threat to small, isolated tiger populations. In Russia, CDV was associated with the deaths of tigers in 2004 and 2010, and was coincident with a localized decline of tigers in Sikhote-Alin Biosphere Zapovednik (from 25 tigers in 2008 to 9 in 2012). Habitat continuity with surrounding areas likely played an important role in promoting an ongoing recovery. We recommend steps be taken to assess the presence and the impact of CDV in all tiger range states, but should not detract focus away from the primary threats to tigers, which include habitat loss and fragmentation, poaching and retaliatory killing. Research priorities include: (i) recognition and diagnosis of clinical cases of CDV in tigers when they occur; and (ii) collection of baseline data on the health of wild tigers. CDV infection of individual tigers need not imply a conservation threat, and modeling should complement disease surveillance and targeted research to assess the potential impact to tiger populations across the range of ecosystems, population densities and climate extremes occupied by tigers. Describing the role of domestic and wild carnivores as contributors to a local CDV reservoir is an important precursor to considering control measures.

New hope for the survival of the Amur leopard in China.

Natural range loss limits the population growth of Asian big cats and may determine their survival. Over the past decade, we collected occurrence data of the critically endangered Amur leopard worldwide and developed a distribution model of the leopard's historical range in northeastern China over the past decade. We were interested to explore how much current range area exists, learn what factors limit their spatial distribution, determine the population size and estimate the extent of potential habitat. Our results identify 48,252 km(2) of current range and 21,173.7 km(2) of suitable habitat patches and these patches may support 195.1 individuals. We found that prey presence drives leopard distribution, that leopard density exhibits a negative response to tiger occurrence and that the largest habitat patch connects with 5,200 km(2)of Russian current range. These insights provide a deeper understanding of the means by which endangered predators might be saved and survival prospects for the Amur leopard not only in China, but also through imperative conservation cooperation internationally.

Social structure and space use of Amur tigers (Panthera tigris altaica) in Southern Russian Far East based on GPS telemetry data.

To better understand the spatial structure of Amur tigers (Panthera tigris altaica) at the southern edge of their range we fitted 14 tigers (6♀♀ and 8♂♂) with 15 GPS-Argos collars between 2008 and 2011 in 2 study sites: the Ussuriskii Reserve of southern Sikhote-Alin and the Land of the Leopard National Park in southwest Primorye, Russian Far East. Fixed kernel estimates of male home ranges were larger than those of female home ranges (P < 0.05 [mean 95% fixed kernel(♀) = 401 ± 205 km(2) ; mean 95% fixed kernel(♂) = 778 ± 267 km(2)]). The home range size of females varied greatly, but on average was similar to estimates derived from earlier work further north. Low overlap of adjacent home ranges suggested that females retained exclusive territories. Real core areas of females overlapped only slightly, and remained stable over multiple years. The home ranges of adult males were smaller than those of males to the north, and in contrast to previous studies, high overlap among males indicated the absence of territoriality. Nonetheless, real core areas of males did not overlap, suggesting some spatial separation. In comparison to other tiger populations and other areas of the Russian Far East, the sex ratio in our 2 study areas was highly skewed towards males. We believe this skewed sex ratio resulted in the dissolution of territoriality of males due to an inability to defend individual females, with males resorting to scramble competition for mates. Continued monitoring of these sites to determine whether shifts in the sex ratio might result in a return to male territoriality would provide confirmation of our tentative hypothesis.

Effects of Roads and Human Disturbance on Amur Tigers.

Understanding the effects of human disturbance on endangered wildlife populations is critical to their conservation. We examined the effects of roads and human disturbance on the survivorship and foraging efficiency of Amur tigers ( Panthera tigris altaica) on and near Sikhote-Alin State Biosphere Zapovednik, Primorye Krai ( province), Russia. To evaluate the effects of roads, we estimated survivorship of radiocollared tigers and their cubs living in three types of areas: (1) areas with primary roads, (2) areas containing secondary roads, and (3) areas with minimal or no road access. We classified a tiger into one of these three treatments based on which types of roads bisected their 50% minimum convex polygon home ranges. Over a 9-year period (1991-2000), adult female survival was greatest (χ 2 = 12.2, df = 2, p = 0.002) for radiocollared tigers in roadless areas. All adult female tigers in roadless areas survived their tenure in those locations (n = 2), whereas all died or disappeared prematurely from areas with primary roads (n = 6). Cub survival was lower in areas with primary and secondary roads than in roadless areas (χ 2 = 10.9, df = 1, p < 0.009). We evaluated the effects of human disturbance at kill sites by examining 86 kills made by 15 tigers determining whether human disturbance had occurred at the kill site, and examining prey carcasses after tigers left, to estimate the percent meat eaten and whether the tiger abandoned the kill following human disturbance. Tigers undisturbed at kills consumed more meat ( Z = 3.71, p = 0.0002) from each kill than disturbed tigers did. Undisturbed tigers also spent more time at each kill site than disturbed tigers did ( Z = 2.3; p = 0.02). Abandonment of kills occurred in 63% of 24 instances when tigers were disturbed by people. Because roads decrease the survivorship and reproductive success of tigers, we recommend that in habitats managed for tigers, construction of new roads should be prohibited wherever possible and access to secondary roads (e.g., logging roads) should be reduced or prevented wherever possible. Protected areas seem to cease functioning as source populations where road access exists, and unprotected areas-the majority of Amur tiger range-cannot sustain stable populations with the increasing threat of human access to tiger habitat.

RESUMEN: Entender los efectos de la perturbación humana sobre poblaciones de vida silvestre en peligro es crítico para su conservación. Examinamos los efectos de caminos y perturbación humana sobre la supervivencia y eficiencia de forrajeo de tigres Amur ( Panthera tigris altaica) en y cerca de Biosfera Estatal Zapovednik Sikhote-Alin, Primorye Krai ( provincia), Rusia. Para evaluar los efectos de los caminos, estimamos la supervivencia de tigres con radio-collares y sus crías en áreas de tres tipos: 1) áreas con caminos primarios, 2) áreas con caminos secundarios y 3) áreas con mínimo o sin acceso de caminos. Clasificamos un tigre en uno de estos tres tratamientos con base en cuales tipos de caminos dividen el 50 % mínimo del polígono convexo de sus rangos de hogar. En un período de 9 años (1991-2000), la supervivencia de hembras adultas fue mayor (χ 2 = 12.2, g.l. = 2, p = 0.002) en tigres con radio-collar en áreas sin caminos. Todas las hembras adultas en áreas sin caminos sobrevivieron su dominio en esas localidades (n = 2), mientras que todas murieron o desaparecieron prematuramente en las áreas con caminos primarios (n = 6). La supervivencia de crías fue menor en áreas con caminos primarios y secundarios que en áreas sin caminos (χ 2 = 10.9, g.l. = 1, p < 0.009). Evaluamos los efectos de la perturbación humana en sitios de depredación mediante el examen de 86 muertes causadas por 15 tigres, determinando si había ocurrido perturbación humana en el sitio, y mediante el examen de restos de las presas después de que se alejaron los tigres, para estimar el porcentaje de carne consumida y si el tigre abandonó su presa después de una perturbación humana. Los tigres no perturbados consumieron más carne ( Z = 3.71, p = 0.0002) de cada presa que los tigres perturbados. Tigres no perturbados pasaron más tiempo con su presa que tigres perturbados ( Z = 2.3, p = 0.02). El abandono de presas ocurrió en 63 % de 24 eventos cuando los tigres fueron perturbados por humanos. Debido a que los caminos reducen la supervivencia y el éxito reproductivo de los tigres, recomendamos que en los hábitats bajo manejo para tigres, se debe prohibir la construcción de caminos donde sea posible y se debe reducir o prevenir el acceso a caminos secundarios (e. g. caminos madereros) donde sea posible. Las áreas protegidas dejan de funcionar como poblaciones fuente donde existe acceso a caminos, y las áreas no protegidas ( la mayor parte del rango de tigre Amur) no pueden sostener poblaciones estables con la incremento en la amenaza del acceso humano al hábitat de los tigres.