Seasonal variation in daily activity patterns of snow leopards and their prey.

The daily and seasonal activity patterns of snow leopards (Panthera uncia) are poorly understood, limiting our ecological understanding and hampering our ability to mitigate threats such as climate change and retaliatory killing in response to livestock predation. We fitted GPS-collars with activity loggers to snow leopards, Siberian ibex (Capra sibirica: their main prey), and domestic goats (Capra hircus: common livestock prey) in Mongolia between 2009 and 2020. Snow leopards were facultatively nocturnal with season-specific crepuscular activity peaks: seasonal activity shifted towards night-sunrise during summer, and day-sunset in winter. Snow leopard activity was in contrast to their prey, which were consistently diurnal. We interpret these results in relation to: (1) darkness as concealment for snow leopards when stalking in an open landscape (nocturnal activity), (2) low-intermediate light preferred for predatory ambush in steep rocky terrain (dawn and dusk activity), and (3) seasonal activity adjustments to facilitate thermoregulation in an extreme environment. These patterns suggest that to minimise human-wildlife conflict, livestock should be corralled at night and dawn in summer, and dusk in winter. It is likely that climate change will intensify seasonal effects on the snow leopard's daily temporal niche for thermoregulation in the future.

Using interview surveys and multispecies occupancy models to inform vertebrate conservation.

Species distribution data are an essential biodiversity variable requiring robust monitoring to inform wildlife conservation. Yet, such data remain inherently sparse because of the logistical challenges of monitoring biodiversity across broad geographic extents. Surveys of people knowledgeable about the occurrence of wildlife provide an opportunity to evaluate species distributions and the ecology of wildlife communities across large spatial scales. We analyzed detection histories of 30 vertebrate species across the Western Ghats biodiversity hotspot in India, obtained from a large-scale interview survey of 2318 people who live and work in the forests of this region. We developed a multispecies occupancy model that simultaneously corrected for false-negative (non-detection) and false-positive (misidentification) errors that interview surveys can be prone to. Using this model, we integrated data across species in composite analyses of the responses of functional species groups (based on disturbance tolerance, diet, and body mass traits) to spatial variation in environmental variables, protection, and anthropogenic pressures. We observed a positive association between forest cover and the occurrence of species with low tolerance of human disturbance. Protected areas were associated with higher occurrence for species across different functional groups compared with unprotected lands. We also observed the occurrence of species with low disturbance tolerance, herbivores, and large-bodied species was negatively associated with developmental pressures, such as human settlements, energy production and mining, and demographic pressures, such as biological resource extraction. For the conservation of threatened vertebrates, our work underscores the importance of maintaining forest cover and reducing deforestation within and outside protected areas, respectively. In addition, mitigating a suite of pervasive human pressures is also crucial for wildlife conservation in one of the world's most densely populated biodiversity hotspots.

Uso de Encuestas y Modelos de Ocupación Multiespecies para Orientar la Conservación de Vertebrados Resumen Los datos de distribución de especies son una variable esencial de la biodiversidad que requieren de un monitoreo sólido para orientar la conservación de la fauna. Aun así, dichos datos permanecen inherentemente escasos debido a los obstáculos logísticos del monitoreo de la biodiversidad a lo largo de extensiones geográficas generalizadas. Las encuestas realizadas a personas conocedoras de la incidencia de fauna proporcionan una oportunidad para evaluar la distribución de las especies y la ecología de las comunidades de fauna en escalas espaciales grandes. Analizamos las historias de detección de 30 especies de vertebrados en los Ghats Occidentales de la India obtenidos a partir de una encuesta a gran escala realizada por entrevistas a 2318 personas que viven y trabajan en los bosques de esta región. Desarrollamos un modelo de ocupación multiespecies que corrigió simultáneamente los errores falsos negativos (no detección) y los falsos positivos (identificación correcta) a los que están propensos las encuestas por entrevista. Con este modelo, integramos los datos de todas las especies a un análisis compuesto de las respuestas de los grupos funcionales de especies (con base en la tolerancia a la perturbación, dieta y características de masa corporal) para la variación espacial en las variables ambientales, protección y presiones antropogénicas. Observamos una asociación positiva entre la incidencia de especies con la baja tolerancia a la perturbación humana y a la cobertura forestal. Las áreas protegidas estuvieron asociadas con una incidencia mayor para las especies ubicadas en diferentes grupos funcionales comparadas con las áreas desprotegidas. También observamos que la incidencia de especies con una tolerancia baja a las perturbaciones, herbívoros y especies de mayor tamaño estaba asociada negativamente con las presiones de desarrollo, como los asentamientos humanos, la producción de energías y minería, y las presiones demográficas, como la extracción de recursos biológicos. Para la conservación de vertebrados amenazados, nuestro trabajo hace hincapié en la importancia de mantener la cobertura forestal y reducir la deforestación dentro y fuera de las áreas protegidas, respectivamente. Además, la mitigación de un conjunto de presiones humanas dominantes también es crucial para la conservación de la naturaleza en uno de los puntos calientes de biodiversidad con una de las mayores densidades poblacionales del mundo.

Increasing risks for emerging infectious diseases within a rapidly changing High Asia.

The cold and arid mountains and plateaus of High Asia, inhabited by a relatively sparse human population, a high density of livestock, and wildlife such as the iconic snow leopard Panthera uncia, are usually considered low risk for disease outbreaks. However, based on current knowledge about drivers of disease emergence, we show that High Asia is rapidly developing conditions that favor increased emergence of infectious diseases and zoonoses. This is because of the existing prevalence of potentially serious pathogens in the system; intensifying environmental degradation; rapid changes in local ecological, socio-ecological, and socio-economic factors; and global risk intensifiers such as climate change and globalization. To better understand and manage the risks posed by diseases to humans, livestock, and wildlife, there is an urgent need for establishing a disease surveillance system and improving human and animal health care. Public health must be integrated with conservation programs, more ecologically sustainable development efforts and long-term disease surveillance.

Spatial variation in population-density of snow leopards in a multiple use landscape in Spiti Valley, Trans-Himalaya.

The endangered snow leopard Panthera uncia occurs in human use landscapes in the mountains of South and Central Asia. Conservationists generally agree that snow leopards must be conserved through a land-sharing approach, rather than land-sparing in the form of strictly protected areas. Effective conservation through land-sharing requires a good understanding of how snow leopards respond to human use of the landscape. Snow leopard density is expected to show spatial variation within a landscape because of variation in the intensity of human use and the quality of habitat. However, snow leopards have been difficult to enumerate and monitor. Variation in the density of snow leopards remains undocumented, and the impact of human use on their populations is poorly understood. We examined spatial variation in snow leopard density in Spiti Valley, an important snow leopard landscape in India, via spatially explicit capture-recapture analysis of camera trap data. We camera trapped an area encompassing a minimum convex polygon of 953 km2. Our best model estimated an overall density of 0.5 (95% CI: 0.31-0.82) mature snow leopards per 100 km2. Using AIC, our best model showed the density of snow leopards to depend on estimated wild prey density, movement about activity centres to depend on altitude, and the expected number of encounters at the activity centre to depend on topography. Models that also used livestock biomass as a density covariate ranked second, but the effect of livestock was weak. Our results highlight the importance of maintaining high density pockets of wild prey populations in multiple-use landscapes to enhance snow leopard conservation.

Assessing the Effectiveness of a Community-based Livestock Insurance Program.

Financial mechanisms to mitigate the costs of negative human-carnivore interactions are frequently promoted to support human coexistence with carnivores. Yet, evidence to support their performance in different settings is scarce. We evaluated a community-based livestock insurance program implemented as part of a broader snow leopard conservation effort in the Tost Tosonbumba Nature Reserve, South Gobi, Mongolia. We assessed program efficiency and effectiveness for snow leopard conservation using a results-based evaluation approach. Data sources included program records from 2009 to 2018, as well as surveys conducted in 2016 and 2017, which allowed us to compare key indicators across communities that participated in the insurance program and control communities. Program coverage and number of livestock insured rapidly increased over the years to reach 65% of households and close to 11,000 livestock. Participants expressed satisfaction with the program and their contributions increased over time, with an increasing proportion (reaching 64% in 2018) originating from participant premiums, suggesting strong community ownership of the program. Participants were less likely to report the intention to kill a snow leopard and reported fewer livestock losses than respondents from control communities, suggesting increased engagement in conservation efforts. These results together suggest that the insurance program achieved its expected objectives, although it is challenging to disentangle the contributions of each individual conservation intervention implemented in intervention communities. However, in the first three years of the program, snow leopard mortalities continued to be reported suggesting that additional interventions were needed to reach impact in terms of reducing retaliatory killings of large carnivores.

Relative influence of wild prey and livestock abundance on carnivore-caused livestock predation.

Conservation conflict over livestock depredation is one of the key drivers of large mammalian carnivore declines worldwide. Mitigating this conflict requires strategies informed by reliable knowledge of factors influencing livestock depredation. Wild prey and livestock abundance are critical factors influencing the extent of livestock depredation. We compared whether the extent of livestock predation by snow leopards Panthera uncia differed in relation to densities of wild prey, livestock, and snow leopards at two sites in Shey Phoksundo National Park, Nepal. We used camera trap-based spatially explicit capture-recapture models to estimate snow leopard density; double-observer surveys to estimate the density of their main prey species, the blue sheep Pseudois nayaur; and interview-based household surveys to estimate livestock population and number of livestock killed by snow leopards. The proportion of livestock lost per household was seven times higher in Upper Dolpa, the site which had higher snow leopard density (2.51 snow leopards per 100 km2) and higher livestock density (17.21 livestock per km2) compared to Lower Dolpa (1.21 snow leopards per 100 km2; 4.5 livestock per km2). The wild prey density was similar across the two sites (1.81 and 1.57 animals per km2 in Upper and Lower Dolpa, respectively). Our results suggest that livestock depredation level may largely be determined by the abundances of the snow leopards and livestock and predation levels on livestock can vary even at similar levels of wild prey density. In large parts of the snow leopard range, livestock production is indispensable to local livelihoods and livestock population is expected to increase to meet the demand of cashmere. Hence, we recommend that any efforts to increase livestock populations or conservation initiatives aimed at recovering or increasing snow leopard population be accompanied by better herding practices (e.g., predator-proof corrals) to protect livestock from snow leopard.

Identification errors in camera-trap studies result in systematic population overestimation.

Reliable assessments of animal abundance are key for successful conservation of endangered species. For elusive animals with individually-unique markings, camera-trap surveys are a benchmark standard for estimating local and global population abundance. Central to the reliability of resulting abundance estimates is the assumption that individuals are accurately identified from photographic captures. To quantify the risk of individual misidentification and its impact on population abundance estimates we performed an experiment under controlled conditions in which 16 captive snow leopards (Panthera uncia) were camera-trapped on 40 occasions and eight observers independently identified individuals and recaptures. Observers misclassified 12.5% of all capture occasions, resulting in systematically inflated population abundance estimates on average by one third (mean ± SD = 35 ± 21%). Our results show that identifying individually-unique individuals from camera-trap photos may not be as reliable as previously believed, implying that elusive and endangered species could be less abundant than current estimates indicate.

Health and zoonotic Infections of snow leopards Panthera unica in the South Gobi desert of Mongolia.

Background: Snow leopards, Panthera uncia, are a threatened apex predator, scattered across the mountains of Central and South Asia. Disease threats to wild snow leopards have not been investigated.Methods and Results: Between 2008 and 2015, twenty snow leopards in the South Gobi desert of Mongolia were captured and immobilised for health screening and radio-collaring. Blood samples and external parasites were collected for pathogen analyses using enzyme-linked immunosorbent assay (ELISA), microscopic agglutination test (MAT), and next-generation sequencing (NGS) techniques. The animals showed no clinical signs of disease, however, serum antibodies to significant zoonotic pathogens were detected. These pathogens included, Coxiella burnetii, (25% prevalence), Leptospira spp., (20%), and Toxoplasma gondii (20%). Ticks collected from snow leopards contained potentially zoonotic bacteria from the genera Bacillus, Bacteroides, Campylobacter, Coxiella, Rickettsia, Staphylococcus and Streptococcus.Conclusions: The zoonotic pathogens identified in this study, in the short-term did not appear to cause illness in the snow leopards, but have caused illness in other wild felids. Therefore, surveillance for pathogens should be implemented to monitor for potential longer- term disease impacts on this snow leopard population.

Impact of wild prey availability on livestock predation by snow leopards.

An increasing proportion of the world's poor is rearing livestock today, and the global livestock population is growing. Livestock predation by large carnivores and their retaliatory killing is becoming an economic and conservation concern. A common recommendation for carnivore conservation and for reducing predation on livestock is to increase wild prey populations based on the assumption that the carnivores will consume this alternative food. Livestock predation, however, could either reduce or intensify with increases in wild prey depending on prey choice and trends in carnivore abundance. We show that the extent of livestock predation by the endangered snow leopard Panthera uncia intensifies with increases in the density of wild ungulate prey, and subsequently stabilizes. We found that snow leopard density, estimated at seven sites, was a positive linear function of the density of wild ungulates-the preferred prey-and showed no discernible relationship with livestock density. We also found that modelled livestock predation increased with livestock density. Our results suggest that snow leopard conservation would benefit from an increase in wild ungulates, but that would intensify the problem of livestock predation for pastoralists. The potential benefits of increased wild prey abundance in reducing livestock predation can be overwhelmed by a resultant increase in snow leopard populations. Snow leopard conservation efforts aimed at facilitating increases in wild prey must be accompanied by greater assistance for better livestock protection and offsetting the economic damage caused by carnivores.

Multiscale factors affecting human attitudes toward snow leopards and wolves.

The threat posed by large carnivores to livestock and humans makes peaceful coexistence between them difficult. Effective implementation of conservation laws and policies depends on the attitudes of local residents toward the target species. There are many known correlates of human attitudes toward carnivores, but they have only been assessed at the scale of the individual. Because human societies are organized hierarchically, attitudes are presumably influenced by different factors at different scales of social organization, but this scale dependence has not been examined. We used structured interview surveys to quantitatively assess the attitudes of a Buddhist pastoral community toward snow leopards (Panthera uncia) and wolves (Canis lupus). We interviewed 381 individuals from 24 villages within 6 study sites across the high-elevation Spiti Valley in the Indian Trans-Himalaya. We gathered information on key explanatory variables that together captured variation in individual and village-level socioeconomic factors. We used hierarchical linear models to examine how the effect of these factors on human attitudes changed with the scale of analysis from the individual to the community. Factors significant at the individual level were gender, education, and age of the respondent (for wolves and snow leopards), number of income sources in the family (wolves), agricultural production, and large-bodied livestock holdings (snow leopards). At the community level, the significant factors included the number of smaller-bodied herded livestock killed by wolves and mean agricultural production (wolves) and village size and large livestock holdings (snow leopards). Our results show that scaling up from the individual to higher levels of social organization can highlight important factors that influence attitudes of people toward wildlife and toward formal conservation efforts in general. Such scale-specific information can help managers apply conservation measures at appropriate scales. Our results reiterate the need for conflict management programs to be multipronged.

Vigorous dynamics underlie a stable population of the endangered snow leopard Panthera uncia in Tost Mountains, South Gobi, Mongolia.

Population monitoring programmes and estimation of vital rates are key to understanding the mechanisms of population growth, decline or stability, and are important for effective conservation action. We report, for the first time, the population trends and vital rates of the endangered snow leopard based on camera trapping over four years in the Tost Mountains, South Gobi, Mongolia. We used robust design multi-season mark-recapture analysis to estimate the trends in abundance, sex ratio, survival probability and the probability of temporary emigration and immigration for adult and young snow leopards. The snow leopard population remained constant over most of the study period, with no apparent growth (λ = 1.08+-0.25). Comparison of model results with the "known population" of radio-collared snow leopards suggested high accuracy in our estimates. Although seemingly stable, vigorous underlying dynamics were evident in this population, with the adult sex ratio shifting from being male-biased to female-biased (1.67 to 0.38 males per female) during the study. Adult survival probability was 0.82 (SE+-0.08) and that of young was 0.83 (SE+-0.15) and 0.77 (SE +-0.2) respectively, before and after the age of 2 years. Young snow leopards showed a high probability of temporary emigration and immigration (0.6, SE +-0.19 and 0.68, SE +-0.32 before and after the age of 2 years) though not the adults (0.02 SE+-0.07). While the current female-bias in the population and the number of cubs born each year seemingly render the study population safe, the vigorous dynamics suggests that the situation can change quickly. The reduction in the proportion of male snow leopards may be indicative of continuing anthropogenic pressures. Our work reiterates the importance of monitoring both the abundance and population dynamics of species for effective conservation.

Role of Tibetan Buddhist monasteries in snow leopard conservation.

The snow leopard (Panthera uncia) inhabits the rugged mountains in 12 countries of Central Asia, including the Tibetan Plateau. Due to poaching, decreased abundance of prey, and habitat degradation, it was listed as endangered by the International Union for Conservation of Nature in 1972. Current conservation strategies, including nature reserves and incentive programs, have limited capacities to protect snow leopards. We investigated the role of Tibetan Buddhist monasteries in snow leopard conservation in the Sanjiangyuan region in China's Qinghai Province on the Tibetan Plateau. From 2009 to 2011, we systematically surveyed snow leopards in the Sanjiangyuan region. We used the MaxEnt model to determine the relation of their presence to environmental variables (e.g., elevation, ruggedness) and to predict snow leopard distribution. Model results showed 89,602 km(2) of snow leopard habitat in the Sanjiangyuan region, of which 7674 km(2) lay within Sanjiangyuan Nature Reserve's core zones. We analyzed the spatial relation between snow leopard habitat and Buddhist monasteries and found that 46% of monasteries were located in snow leopard habitat and 90% were within 5 km of snow leopard habitat. The 336 monasteries in the Sanjiangyuan region could protect more snow leopard habitat (8342 km(2) ) through social norms and active patrols than the nature reserve's core zones. We conducted 144 household interviews to identify local herders' attitudes and behavior toward snow leopards and other wildlife. Most local herders claimed that they did not kill wildlife, and 42% said they did not kill wildlife because it was a sin in Buddhism. Our results indicate monasteries play an important role in snow leopard conservation. Monastery-based snow leopard conservation could be extended to other Tibetan Buddhist regions that in total would encompass about 80% of the global range of snow leopards.

Globalization of the cashmere market and the decline of large mammals in central Asia.

As drivers of terrestrial ecosystems, humans have replaced large carnivores in most areas, and human influence not only exerts striking ecological pressures on biodiversity at local scales but also has indirect effects in distant corners of the world. We suggest that the multibillion dollar cashmere industry creates economic motivations that link western fashion preferences for cashmere to land use in Central Asia. This penchant for stylish clothing, in turn, encourages herders to increase livestock production which affects persistence of over 6 endangered large mammals in these remote, arid ecosystems. We hypothesized that global trade in cashmere has strong negative effects on native large mammals of deserts and grassland where cashmere-producing goats are raised. We used time series data, ecological snapshots of the biomass of native and domestic ungulates, and ecologically and behaviorally based fieldwork to test our hypothesis. In Mongolia increases in domestic goat production were associated with a 3-fold increase in local profits for herders coexisting with endangered saiga (Saiga tatarica).That increasing domestic grazing pressure carries fitness consequences was inferred on the basis of an approximately 4-fold difference in juvenile recruitment among blue sheep (Pseudois nayaur)in trans-Himalayan India. Across 7 study areas in Mongolia, India, and China's Tibetan Plateau, native ungulate biomass is now <5% that of domestic species. Such trends suggest ecosystem degradation and decreased capacity for the persistence of native species, including at least 8 Asian endemic species: saiga, chiru (Pantholops hodgsoni), Bactrian camel (Camelus bactrianus), snow leopard(Panthera uncia), khulan(Equus hemionus), kiang (E. kiang), takhi (E. przewalski), and wild yak (Bos mutus). Our results suggest striking yet indirect and unintended actions that link trophic-level effects to markets induced by the trade for cashmere.

Reversible immobilization of free-ranging snow leopards (panthera uncia) with a combination of medetomidine and tiletamine-zolazepam.

Conservation and research of the elusive snow leopard (Panthera uncia) have been hampered by inadequate knowledge about its basic life history. Global positioning system (GPS) collars can provide useful information, but there has been limited information available on safe capture methods, drug doses, and efficacy for effective immobilization of free-ranging snow leopards. We describe a drug protocol using a combination of medetomidine and tiletamine-zolazepam for the chemical immobilization of free-ranging snow leopards. We also describe physiologic responses to immobilization drugs, including rectal temperature, heart rate, respiratory rate, and relative hemoglobin oxygen saturation (SpO2) recorded every 10 min. Our study was carried out in the Tost Mountains adjacent to the Great Gobi Desert, in southern Mongolia, between August 2008 and April 2012. Eighteen snow leopards were captured or recaptured with foot-snares on 42 occasions and anesthetized for marking with GPS collars. The snow leopards received on average (±SD) 0.020±0.04 mg/kg body mass medetomidine and 2.17±0.45 mg/kg tiletamine-zolazepam. The duration of ensuing anesthesia was 69±13 min, including an induction period of 10 (±4) min. Anesthesia was reversed with 4 mg (0.10±0.04 mg/kg) atipamezole administered intramuscularly. The mean value for SpO2 for the 37 captures where we could record physiologic values was 91±4. The SpO2 increased significantly during anesthesia (+0.06±0.02%/min), whereas rectal temperature (average 38.1±0.7 C/min, change -0.04±0.003 C/min), heart rate (average 97±9 beats/min, change -0.20±0.03 beats/min), and respiratory rate (average 26±6 breaths/min, change -0.11±0.03 breaths/min) decreased significantly. A dose of 80 mg tiletamine-zolazepam (2 mg/kg body weight) and 0.72 mg medetomidine (0.02 mg/kg body weight) safely immobilized all adult and subadult snow leopards (weight 25-45 kg) in our study. All measured physiologic values remained within clinically acceptable limits.

Standardizing the double-observer survey method for estimating mountain ungulate prey of the endangered snow leopard.

Mountain ungulates around the world have been threatened by illegal hunting, habitat modification, increased livestock grazing, disease and development. Mountain ungulates play an important functional role in grasslands as primary consumers and as prey for wild carnivores, and monitoring of their populations is important for conservation purposes. However, most of the several currently available methods of estimating wild ungulate abundance are either difficult to implement or too expensive for mountainous terrain. A rigorous method of sampling ungulate abundance in mountainous areas that can allow for some measure of sampling error is therefore much needed. To this end, we used a combination of field data and computer simulations to test the critical assumptions associated with double-observer technique based on capture-recapture theory. The technique was modified and adapted to estimate the populations of bharal (Pseudois nayaur) and ibex (Capra sibirica) at five different sites. Conducting the two double-observer surveys simultaneously led to underestimation of the population by 15%. We therefore recommend separating the surveys in space or time. The overall detection probability for the two observers was 0.74 and 0.79. Our surveys estimated mountain ungulate populations (± 95% confidence interval) of 735 (± 44), 580 (± 46), 509 (± 53), 184 (± 40) and 30 (± 14) individuals at the five sites, respectively. A detection probability of 0.75 was found to be sufficient to detect a change of 20% in populations of >420 individuals. Based on these results, we believe that this method is sufficiently precise for scientific and conservation purposes and therefore recommend the use of the double-observer approach (with the two surveys separated in time or space) for the estimation and monitoring of mountain ungulate populations.

Biomass and diversity of dry alpine plant communities along altitudinal gradients in the Himalayas.

A non-linear relationship between phytodiversity and altitude has widely been reported, but the relationship between phytomass and altitude remains little understood. We examined the phytomass and diversity of vascular plants along altitudinal gradients on the dry alpine rangelands of Ladakh, western Himalaya. We used generalized linear and generalized additive models to assess the relationship between these vegetation parameters and altitude. We found a hump-shaped relationship between aboveground phytomass and altitude. We suspect that this is engendered by low rainfall and trampling/excessive grazing at lower slopes by domestic livestock, and low temperature and low nutrient levels at higher slopes. We also found a unimodal relationship between plant species-richness and altitude at a single mountain as well as at the scale of entire Ladakh. The species-richness at the single mountain peaked between 5,000 and 5,200 m, while it peaked between 3,500 and 4,000 m at entire Ladakh level. Perhaps biotic factors such as grazing and precipitation are, respectively, important in generating this pattern at the single mountain and entire Ladakh.

Why should a grazer browse? Livestock impact on winter resource use by bharal Pseudois nayaur.

Many mammalian herbivores show a temporal diet variation between graminoid-dominated and browse-dominated diets. We determined the causes of such a diet shift and its implications for conservation of a medium-sized ungulate-the bharal Pseudois nayaur. Past studies show that the bharal diet is dominated by graminoids (>80%) during summer, but the contribution of graminoids declines to about 50% in winter. We tested the predictions generated by two alternative hypotheses explaining the decline: low graminoid availability during winter causes bharal to include browse in their diet; bharal include browse, with relatively higher nutritional quality, in their diet to compensate for the poor quality of graminoids during winter. We measured winter graminoid availability in areas with no livestock grazing, areas with relatively moderate livestock grazing, and those with intense livestock grazing pressures. The chemical composition of plants contributing to the bharal diet was analysed. The bharal diet was quantified through signs of feeding on vegetation at feeding locations. Population structures of bharal populations were recorded using a total count method. Graminoid availability was highest in areas without livestock grazing, followed by areas with moderate and intense livestock grazing. The bharal diet was dominated by graminoids (73%) in areas with highest graminoid availability. Graminoid contribution to the bharal diet declined monotonically (50, 36%) with a decline in graminoid availability. Bharal young to female ratio was 3 times higher in areas with high graminoid availability than areas with low graminoid availability. The composition of the bharal winter diet was governed predominantly by the availability of graminoids in the rangelands. Our results suggest that bharal include more browse in their diet during winter due to competition from livestock for graminoids. Since livestock grazing reduces graminoid availability, creation of livestock-free areas is necessary for the conservation of grazing species such as the bharal and its predators including the endangered snow leopard in the Trans-Himalaya.

Winter ecology of the Arunachal macaque Macaca munzala in Pangchen Valley, western Arunachal Pradesh, northeastern India.

The newly described Arunachal macaque Macaca munzala occurs largely in sub-tropical to temperate environments at elevations of c. 1,800-3,000 m in Arunachal Pradesh, northeastern India. We studied its over-wintering strategy by comparing the diet, ranging, and behavior of a troop of 24 individuals during winter and spring (December 2005 to May 2006) through instantaneous scan sampling (3,002 records, 448 scans, 112 hr of observation). We also monitored the phenology of food plants. The macaques spent more time (41-66%) feeding in the winter than in spring (33-51%), whereas time spent moving and resting was greater in spring. The diet composed largely of plants, with animal matter being eaten rarely. The number of plant species in the diet increased from 18 to 25 whereas food types rose from 18 to 36 from winter to spring, respectively. Although only two species formed 75% of the winter diet, seven species comprised this proportion in spring. Availability of fruits and young leaves increased in spring; the troop moved more and utilized a larger part of its range during this time. Seasonal changes in behavior could be explained by the scarcity of food and the costs of thermoregulation in winter. Our study suggests that the Arunachal macaque inhabits a highly seasonal environment and has an over-wintering strategy that includes subsisting on a high-fiber diet by increasing the time spent feeding, and minimizing energy expenditure by reducing the time spent moving.