Abstract: Foreword: Field monitoring of wild animal species is rarely accomplished without challenges. Logistical, environmental, and ecological factors dictate the need for appropriate sampling regardless of location, taxa, or objectives. With regards to felids there is no questioning their popularity when it comes to field research. Their role and impact on the ecosystems combined with their “hypercarnivore” lifestyle and cultural significance have resulted in an extensive and historical resume of field studies across the globe. Despite the vast number of studies on felids, there is a significant skew toward the big cats with the scientific knowledge base for small-bodied cat species, including the Pallas’s cat or manul (Otocolobus manul), much smaller. Given the solitary and elusive nature of most small cats, like the manul, that inhabit remote environments, it is not difficult to understand this gap in field research.
Recognising this gap and following years of field research and conservation, the Pallas’s cat International Conservation Alliance (PICA) and their conservation partners identified the need for targeted and easy to follow guidance on best practices for monitoring the manul in the wild. This guide details an extensive compilation of data collection methods and monitoring techniques for the manul that will help practitioners deliver more effective conservation and research efforts.
This guide was edited by Ehsan Moqanaki and Gustaf Samelius that, in close collaboration with a number of co-authors, have brought together a wealth of knowledge on surveying and monitoring manul populations. Each chapter compiles tried and tested techniques from a range of carnivore research projects over the last few decades, with a focus on the manul. In doing so this guide provides a detailed insight into the most effective data collection methods to enhance future conservation and research efforts for the species. We are grateful to all involved in the development of this book and hope that it serves as a valuable practical guide to current and future conservation and researcher efforts, while contributing to long-term conservation actions for this amazing small cat.
The Pallas’s cat International Conservation Alliance

Abstract: The endangered Snow leopard (Panthera uncia) inhabits the high mountain regions through central Asia and is subjected to numerous threats including poaching for traditional Chinese medicine, retribution killing for preying on domestic stock, and habitat fragmentation. However the occurrence and impact of disease on snow leopard populations is unknown. As emerging infectious diseases of wildlife can be an insidious yet important cause of population decline due to mortality or reproductive failure, my study aimed initially to gain knowledge of pathogens circulating among wild and domestic hosts in this region. I used a broad One Health approach to survey a range of species to collect data on disease occurrence that would be useful in improving human and livestock health, as well as snow leopard conservation.
This study is set in the Tost Mountains of the South Gobi Desert of Mongolia and was prompted due to the unexplained deaths of four snow leopards detected within a short timeframe during an ecological study by members of the Snow Leopard Trust. However, investigating disease occurrence in remote, rare and endangered species is a challenge due to inaccessibility of sites, difficulty of capture, and processing samples without facilities.
A One Health approach uses multidisciplinary expertise such as ecological, medical and veterinary, to understand host, pathogen and environmental disease factors. This approach is especially useful for diseases that transfer between people, domestic animals and wildlife. As snow leopards are a rare and elusive species, my surveys were aimed at assessing pathogens circulating in snow leopards as well as in sympatric wild and domestic animals. I collected samples from the following hosts: snow leopards – the target species; rodents which are ubiquitous over the study area and are a suitable sentinel species; ibex which are a native ungulate and natural prey species of the snow leopard; domestic goats which are also a prey species of the snow leopard; free-ranging domestic dogs which interact with the goats. The local indigenous people interact with all these species including snow leopards, mostly via retribution killing. Water samples were also collected from waterholes and wells, which are communal meeting places as drinking sources for all species, hence enabling pathogen exchange. Samples collected included blood samples, faecal samples or rectal swabs and ectoparasites if present. These samples were transported to laboratories in Sweden and Belgium where I conducted diagnostic assays for zoonotic pathogens that are present in other regions of Mongolia and impact the health of humans and animals. I used enzyme- linked immune assay (ELISA), polymerase chain reaction (PCR) and next-generation sequencing (NGS) for pathogens including Coxiella burnetii, Toxoplasma gondii, Leptospira spp., Brucella spp., Yersinia pestis and tick borne encephalitis virus. Serovars of Leptospira were elucidated using microscopic agglutination tests (MAT). The dog blood samples were also tested for canine distemper virus. Ticks, faeces, rectal swabs and water were tested for bacteria, Echinococcus, Giardia and Cryptosporidium using PCR and NGS.
Health records for humans and animals in the region were not available so, in addition to testing animal samples, I used questionnaire surveys to obtain information on perceptions of the herders concerning health of their families, their domestic animals and wildlife. Questions also assessed preventative health management and treatments used.
Over three field trips I caught and sampled twenty snow leopards, 177 rodents (8 species), 41 dogs and 270 goats. I also sampled 11 waterholes/wells, and preserved 18 ticks, hundreds of fleas and collected faecal samples from ibex.
Most animals that were sampled and examined clinically appeared in good health, but the serosurvey revealed a moderate to high level of exposure to serious pathogens: C. burnetii, T. gondii and Leptospira spp. There were no published reports of human infections with these pathogens in the study area, which is likely due to a lack of testing.
Snow leopards had the highest prevalence of C. burnetii antibodies (25%), followed by rodents (16%), dogs (10%) and goats (9.5%). Goats had the highest prevalence of T. gondii antibodies (90%), dogs (66%), snow leopards (20%) and rodents (16%). Rodents had the highest prevalence of Leptospira spp. (34%), followed by snow leopards (20%) and dogs (5%). Serovars interrogans Australis was identified in the rodents and snow leopards and interrogans Ictohaemorrhagiae was identified in the rodents and dogs. Other serovars were also present from the results of the ELISA but did not match those listed in the MAT panel, so could not be identified. Goats were not tested for infection with leptospirosis. Brucella was not identified in the goats even though it occurs at high prevalence in stock in the rest of Mongolia where it is a large health and economic concern. In rodents, the zoonotic Puumala and Seoul hantavirus were identified for the first time in Mongolia. Analysis of data from rodents showed the pathogens detected (C. burnetii, T.gondii, Hanta virus and Leptospira spp.) differed significantly in prevalence, with a strong year effect driven mainly by Leptospira, which increased in prevalence across the three year study period. Toxoplasma gondii differed slightly in prevalence among rodent species. There was no significant difference in prevalence of interaction of pathogens among years or rodent species.
Poor health was detected in goats with 10 out of the 14 goats tested via haematology and biochemistry being anaemic with haematocrits less than 20%. Haematology and biochemistry values for the other animal species appeared normal. I established haematology and biochemistry reference tables for two rodent species – red-cheeked ground squirrels and jerboas.
Water samples were negative for serious pathogens. Fleas were negative for Yersinia pestis. However, ticks were positive for several genera of potential zoonoses, including Anaplasma, Bacillus, Coxiella, Clostridia, Francisella, Rickettsia, Staphylococcus, Streptococcus and Yersinia. Faecal samples were also positive for genera of potentially zoonotic bacteria including those listed above plus Bacteroides, Bordetella, Campylobacter and Enterococcus.
Results from the two questionnaire surveys revealed the main reported illness in people were colds and flu. However, the local doctor also reported hepatitis as common. She also said that the local people contracted brucellosis whereas I did not identify this pathogen in their livestock. The herders thought their main loss of stock was from predation, with wolves identified as the main predator and snow leopards as the second. Other causes of stock loss perceived as important were adverse climatic conditions such as drought or severe winters while infectious disease was not a concern. Results from these surveys also highlighted gaps in health care for humans and livestock, especially around vaccination and parasite treatments.
In summary, I found that snow leopards and other wild and domestic animals within the study area tested positive for previous exposure to several important zoonotic pathogens. These pathogens were likely circulating among species via contamination of pasture and via predation and have potential to cause illness and reproductive loss. However, I detected no adverse effects on the health of the animals due to infection with these pathogens, and observed no related mortality or illness during my field trips. Hence the deaths of the four snow leopards that were the impetus for my study have not been explained, and monitoring and surveillance of this population should continue.
My findings on wildlife and domestic animal pathogens have relative importance to improving productivity of livestock and the health of the nomadic herders. I recommend improving the health of goats through vaccination and anti-parasite programmes, which will improve their fecundity and survival and thus increase herder income. These programmes will also have flow-on effects to improve the health of the native ungulates that share the grazing areas by decreasing the risk of pathogen transfer between them and also to the snow leopards that prey on them. Demonstrating the importance of herd health may also help mitigate herder wildlife conflict as increased productivity could decrease the perceived importance of predation on herd numbers.
Coxiella burnetii and Leptospires spp are a likely cause of illness in people, despite the lack of reported diagnoses. As rodents had a moderate prevalence of all pathogens tested and inhabit the gers of the local people, it is important to raise awareness of the risk of pathogen transfer to people via rodent excrement contaminating stored food and eating utensils. Risk of human exposure to pathogens during goat slaughter can also be reduced via improved hygiene practices.
By identifying pathogens with broad host ranges in a variety of species in this remote mountainous region, my study provides the basis for understanding health risks to wildlife, domestic animals and humans. Consideration of likely transmission routes for pathogens between species can inform current recommendations to improve health, productivity and hence conservation, of the endangered snow leopard – The Ghost of the Mountain.

Abstract: There is a growing agreement that conservation needs to be proactive and pay increased attention to common species and to the threats they face. The blue sheep (Pseudois nayaur) plays a key ecological role in sensitive high-altitude ecosystems of Central Asia and is among the main prey species for the globally vulnerable snow leopard (Panthera uncia). As the blue sheep has been increasingly exposed to human pressures, it is vital to estimate its population dynamics, protect the key populations, identify important habitats, and secure a balance between conservation and local livelihoods. We conducted a study in Manang, Annapurna Conservation Area (Nepal), to survey blue sheep on 60 transects in spring (127.9 km) and 61 transects in autumn (134.7 km) of 2019, estimate their minimum densities from total counts, compare these densities with previous estimates, and assess blue sheep habitat selection by the application of generalized additive models (GAMs). Total counts yielded minimum density estimates of 6.0–7.7 and 6.9–7.8 individuals/km2 in spring and autumn, respectively, which are relatively high compared to other areas. Elevation and, to a lesser extent, land cover indicated by the normalized difference vegetation index (NDVI) strongly affected habitat selection by blue sheep, whereas the effects of anthropogenic variables were insignificant. Animals were found mainly in habitats associated with grasslands and shrublands at elevations between 4,200 and 4,700 m. We show that the blue sheep population size in Manang has been largely maintained over the past three decades, indicating the success of the integrated conservation and development efforts in this area. Considering a strong dependence of snow leopards on blue sheep, these findings give hope for the long-term conservation of this big cat in Manang. We suggest that long-term population monitoring and a better understanding of blue sheep–livestock interactions are crucial to maintain healthy populations of blue sheep and, as a consequence, of snow leopards.

Abstract: Predators shape their ecosystems through myriad interactions with prey, other predators, and humans. However, the effects of these interactions may be contingent on multiple contextual factors, hindering prediction in any given community and impeding a general understanding of the ecological effects of predators. Despite their prominence as conservation flagship in the mountains of Central Asia, even basic aspects of snow leopard (Panthera uncia) ecology remain underexplored and poorly understood. The ecology of wolves (Canis lupus), sympatric with snow leopards throughout that species’ range, has been even more neglected in the region, notwithstanding the significant impact of livestock depredation on pastoralist communities. This dissertation examines the interactions underlying the coexistence of wolves and snow leopards, including those with humans and their joint effects on prey, with the broader goal of improving our understanding of the context-dependence of the non-consumptive effects (NCEs) of predators. In Chapter 2, I explore the patterns of spatial, temporal, and dietary niche overlap between wolves and snow leopards in the Eastern Pamir Mountains of Tajikistan. I show that in
light of dietary and temporal overlap, the two predators’ coexistence may depend on strong spatial partitioning. In Chapter 3, I explore the consequences of this spatial partitioning by investigating how shared prey with distinct escape tactics, ibex (Capra sibirica) and argali (Ovis ammon), navigate the tradeoffs posed by the two predators in the Central Tien Shan Mountains of Kyrgyzstan. Each ungulate responded to each predator in a manner that was predictable based on the compatibility of their respective evasion and hunting-mode traits, suggesting that non- consumptive predator effects depend not on predator hunting mode or prey escape tactics, but rather on their interaction. Furthermore, short-term predation risk may upend each ungulates’ long-term risk avoidance strategy, suggesting that emergent effects of multiple predators may have important consequences in this system. In Chapter 4, I develop a novel approach to investigate large-scale patterns of livestock depredation risk and occurrence for wolves and snow leopards, but also lynx (Lynx lynx) and bears (Ursus arctos), in the Western Pamirs of Tajikistan. Livestock depredation was commonplace, with most communities exposed to multiple predators, highlighting that conservation efforts meant to reduce conflict between people and carnivores should aim to reduce depredation as it is experienced by human communities – a threat from the entire carnivore guild. Overall, my results suggest that single-species approaches to conservation in the mountains of Central Asia may be inadequate for ecosystems and people. This dissertation advances the cause of conservation in Central Asia by providing an empirical perspective on how snow leopards and wolves coexist and shape their ecosystems, and by providing practical insight into the challenge of livestock depredation and conflict, a primary threat to wolves and snow leopards in the region. By showing that the non-consumptive effects of predators cannot be predicted based solely on prey escape tactics or predator hunting mode alone, it also contributes to a more comprehensive understanding of the role of predators in shaping ecosystems.

Abstract: The snow leopard (Panthera uncia) is anatomically and physiologically adapted for life in the rocky terrain of alpine zones in Central and South Asia. Panthera uncia is scansorial, and typically hunts solitarily by using overhead ambush of prey, rather than the typical stalking pattern of other large pantherines. In this study, we conducted dissections, detailed documentation, and illustrated the forelimb anatomy of two adult P. uncia specimens (1M/1F). Qualitative and quantitative data revealed an intriguing combination of functional adaptations illustrating a balance between the diverse demands of head-first descent, pouncing, climbing across rocky terrain, restraint of large prey, rapid pursuit, and navigating deep snow. In many forelimb proportions, P. uncia is intermediate between the cursorial Acinonyx jubatus (cheetah) and the scansorial forest dwelling Panthera onca (jaguar). Enlarged scapular and pectoral musculature provide stability to the shoulder girdle during grappling with large prey, as well as support during jumping and climbing. A small, unarticulated bony clavicle may provide greater stability to the forelimb, while still allowing flexibility. In the brachium and antebrachium of P. uncia, there is a functional compromise between the powerful grip needed for grasping large prey and the stability necessary for rapid pursuit of prey over uneven, rocky terrain. A unique bifurcation in the tendon of m. biceps brachii may provide additional functional stability at the radiohumeral joint. Intrinsic muscles of the palmar manus are broad and fleshy, acting as an enlarged surface area to evenly distribute body weight while walking on soft snow. However, muscles that act to provide fine manual manipulation are reduced, as in other large prey specialists. Overall, P. uncia displays morphological adaptive parallels with scansorial, large prey spe- cializing pantherines, such as P. onca, while also showing adaptations for running.

Abstract: Large mammalian carnivores have undergone catastrophic declines during the Anthropocene across the world. Despite their pivotal roles as apex predators in food webs and ecosystem dynamics, few detailed di- etary datasets of large carnivores exist, prohibiting deep understanding of their coexistence and persistence in human-dominated landscapes. Here, we present fine-scaled, quantitative trophic interactions among sym- patric carnivores from three assemblages in the Mountains of Southwest China, a global biodiversity hotspot harboring the world’s richest large-carnivore diversity, derived from DNA metabarcoding of 1,097 fecal sam- ples. These assemblages comprise a large-carnivore guild ranging from zero to five species along with two mesocarnivore species. We constructed predator-prey food webs for each assemblage and identified 95 vertebrate prey taxa and 260 feeding interactions in sum. Each carnivore species consumed 6–39 prey taxa, and dietary diversity decreased with increased carnivore body mass across guilds. Dietary partitioning was more evident between large-carnivore and mesocarnivore guilds, yet different large carnivores showed divergent proportional utilization of different-sized prey correlating with their own body masses. Large car- nivores particularly selected livestock in Tibetan-dominated regions, where the indigenous people show high tolerance toward wild predators. Our results suggest that dietary niche partitioning and livestock subsidies facilitate large-carnivore sympatry and persistence and have key implications for sustainable conservation promoting human-carnivore coexistence.

Abstract: Understanding landscape connectivity and population genetic parameters is imperative for threatened species management. However, such information is lacking for the snow leopard (Panthera uncia). This study sought to explore hierarchical snow leopard gene flow patterns and drivers of genetic structure in Mongolia and China. A total of 97 individuals from across Mongolia and from the north-eastern edge of the Qinghai-Tibetan Plateau in Gansu Province to the middle of Qinghai Province in China were genotyped across 24 microsatellite loci. Distance-based frameworks were used to determine a landscape scenario best explaining observed genetic structure. Spatial and non-spatial methods were used to investigate fine-scale autocorrelation and similarity patterns as well as genetic structure and admixture. A genetic macro-division between populations in China and Mongolia was observed, suggesting that the Gobi Desert is a substantial barrier to gene flow. However, admixture and support for a resistance-based mode of isolation suggests connective routes that could facilitate movement. Populations in Mongolia had greater connectivity, indicative of more continuous habitat. Drivers of genetic structure in China were difficult to discern, and fine-scale sampling is needed. This study elucidates snow leopard landscape connectivity and helps to prioritize conservation areas. Although contact zones may have existed and occasional crossings can occur, establishing corridors to connect these areas should not be a priority. Focus should be placed on maintaining the relatively high connectivity for snow leopard populations within Mongolia and increasing research efforts in China.

Abstract: The pastoral practices of the Dokpa herders of North Sikkim have been transforming in response to the geo-political and socio-economic changes in the region. Against the backdrop of these changes, this study aims to understand the current state of pastoralism in North Sikkim with three specific objectives: (i) to understand the current rangeland management practices of the Dokpa community; (ii) to examine the social, political and ecological stresses to continuity of traditional pastoral livelihoods; and (iii) to document the baseline on human-wildlife relations. We focused on one of the two subset populations of Dokpa herders of North Sikkim and, using a mixed-methods approach, conducted 12 semi-structured interviews, four key respondent interviews and two focused group discussions. The resource use by the Dokpas is unique, and unlike the rest of the Himalayan range, they access the high-altitude pastures in winters and the lower ones in summer. Pastures in the higher altitudes experience heavier winds, which leads to lower levels of snow deposition — thus ensuring access to dried pasture forage for livestock during the lean season. The decisions pertaining to resource management are taken by the head of the local institution Dzumsa, the Pipon. Primary stresses to the continuation of traditional pastoral practices are fragmentation of pastureland post- Sino-Indian war of 1962 and the consequent establishment of armed forces, livestock depredation by free-ranging dogs followed by wild predators and continued socio-economic marginalisation of the pastoralists under a supposedly egalitarian institutional regime. Extreme climatic events in the recent past have also contributed to significant livestock loss. Dokpa transhumant practices are on an overall decline, with most members of the younger generation shifting to non-herding livelihoods. The availability of alternate livelihood options with the improved connectivity, access to education and development of the tourism industry has led to changing aspirations of the younger generations. In only two of the twelve households we surveyed, the younger generation continues herding, while the rest have moved to the cities and towns. In terms of human-wildlife relations, the respondents mostly hold a positive attitude towards wildlife and conservation actions despite livestock predation by wild predators, since the free-ranging dogs cause the highest livestock loss. With the inputs from the Dokpas, we provide recommendations towards a facilitative environment for the continuation of the traditional herding in the region, which is critical for the survival of pastoralism in North Sikkim, presently hinged on less than two dozen of elderly Dokpas.

Abstract: In the present study, we describe an interspecific kleptoparasitic interaction between two sympatric mammalian carnivores in the high altitudinal Trans-Himalaya region of Himachal Pradesh, India. The study was based on the inferences drawn from the circumstantial evidence (direct and indirect) noticed in the study area in Pin Valley National Park. The inferences from the analysis of the evidence suggested the interaction between a Snow Leopard Panthera uncia, a Red Fox Vulpes vulpes, and a donkey. The arrangement of evidence in a sequential manner suggested that a donkey was killed by a Snow Leopard and a Red Fox stole the food from the carrion of the Snow Leopard’s prey. The Red Fox was killed by the Snow Leopard, which was caught while stealing. The present study represents an example of kleptoparasitic interaction between the Snow Leopard and the Red Fox. This study also proves that such interactions may cost the life of a kleptoparasite and supports the retaliation behaviour of Snow Leopards.

Abstract: 1. Spatial responses to risk from multiple predators can precipitate emergent consequences for prey (i.e. multiple-predator effects, MPEs) and mediate indirect interactions between predators. How prey navigate risk from multiple predators may therefore have important ramifications for understanding the propagation of predation-risk effects (PREs) through ecosystems.
2. The interaction of predator and prey traits has emerged as a potentially key driver of antipredator behaviour but remains underexplored in large vertebrate systems, particularly where sympatric prey share multiple predators. We sought to better generalize our understanding of how predators influence their ecosystems by considering how multiple sources of contingency drive prey distribution in a multi-predator–multi-prey system.
3. Specifically, we explored how two sympatric ungulates with different escape tactics—vertically agile, scrambling ibex Capra sibirica and sprinting argali Ovis ammon—responded to predation risk from shared predators with contrasting hunting modes—cursorial wolves Canis lupus and vertical-ambushing, stalking snow leopards Panthera uncia.
4. Contrasting risk posed by the two predators presented prey with clear trade-offs. Ibex selected for greater exposure to chronic long-term risk from snow leopards, and argali for wolves, in a nearly symmetrical manner that was predictable based on the compatibility of their respective traits. Yet, acute short-term risk from the same predator upended these long-term strategies, increasing each ungulates' exposure to risk from the alternate predator in a manner consistent with a scenario in which conflicting antipredator behaviours precipitate risk-enhancing MPEs and mediate predator facilitation. By contrast, reactive responses to wolves led ibex to reduce their exposure to risk from both predators—a risk-reducing MPE. Evidence of a similar reactive risk-reducing effect for argali vis-à-vis snow leopards was lacking.
5. Our results suggest that prey spatial responses and any resulting MPEs and prey-mediated interactions between predators are contingent on the interplay of hunting mode and escape tactics. Further investigation of interactions among various drivers of contingency in PREs will contribute to a more comprehensive understanding and improved forecasting of the ecological effects of predators.

Abstract: Animal-borne tracking devices have generated a wealth of new knowledge, allowing us to better understand, manage and conserve species. Fitting such tracking devices requires that animals are captured and often chemically immobilized. Such procedures cause stress and involve the risk of injuries and loss of life even in healthy individuals. For telemetry studies to be justifiable, it is vital that capture operations are planned and executed in an efficient and ethical way. Project objectives must be clearly articulated to address well-defined knowledge gaps, and studies designed to maximize the probability of achieving those goals. We provide guidelines for how to plan, design, and implement telemetry studies with a special emphasis on snow leopards that are typically captured using foot snares. We also describe the necessary steps to ensure that captures are conducted safely, and with minimal stress to animals.

Abstract: Human conflict with large carnivores continues to be a great conservation challenge, and conflict with snow leopards (Panthera uncia) has been studied to understand causes and propose mitigation schemes. While the nature of snow leopard-human conflict is similar in most cases, reported studies have been case- and area-specific with mitigation strategies not necessarily based on a synthesis of relevant literature. We reviewed snow leopard literature published from 1970-2020 to identify the main drivers of human-snow leopard conflict (HSLC) and describe conservation and conflict mitigation strategies commonly employed. Based on 47 relevant peer-reviewed articles, review papers, book chapters, project reports, and other grey literature, we identified four major conflict domains: livestock management-related, socio-economic/human-related, ecological, and policy-related. Most articles suggested more than one conflict mitigation scheme. Three conflict mitigation domains – preventive, supportive, and compensatory – were widely reflected in the snow leopard-human conflict literature. The most commonly reported mitigation schemes included: 1) building or predator-proofing corrals; 2) training shepherds and improving livestock guarding; 3) livestock insurance schemes; 4) compensation for livestock predation; 5) capacity building, education, and awareness programs; and 6) improved breeding and use of guard dogs. Future management efforts need to tailor their approach depending on cultural, economic, and ecological circumstances.