Andrew Marten Kittle

As humans and wildlife increasingly overlap on a rapidly changing planet, it is important to understand co-existence mechanisms in unprotected landscapes. This is especially so for large carnivores, with which humans share an uneasy history. In Sri Lanka’s tea estate-dominated Central Highlands, a dense, widespread human population shares the landscape with the island’s apex predator, the Sri Lankan leopard ( Panthera pardus kotiya ). This study used remote cameras to determine leopard density, generalized linear models to investigate leopard space-use in relation to key anthropogenic, ecological and environmental variables, and fitted kernel density curves to identify temporal activity patterns in this unprotected, human-dominated region. Adult leopard density (8.82/100 km ² ) was lower here than in key Sri Lankan National Parks, but not significantly, suggesting good leopard habitat suitability. Spatially, leopards did not avoid humans but incorporated anthropogenic factors into space-use decisions, with areas of intermediate human and dog presence disproportionately used. Male leopards were more likely to occur at intermediate distances from human settlements, perhaps balancing risk with efficiency while navigating this shared landscape. Female leopards biased space-use towards areas of high natural prey availability, aligning with their need to provision cubs. Temporally, leopards were significantly less diurnal and more active at dusk than their protected counterparts. This was likely due to temporal partitioning from diurnally active humans, an adaptation which allows for human-leopard co-existence. This is the first comprehensive study of leopards outside protected areas in Sri Lanka, and highlights the value of unprotected landscapes for long-term leopard conservation. To foster human-leopard co-existence it is necessary to maintain current human activity patterns which allow for temporal partitioning; ensure existing natural prey resources remain to prevent leopards switching to domestic prey species; and reduce/eliminate the use of wire snares which transcend the effect of temporal partitioning by de-coupling persecution risk from human presence.

The endemic Sri Lankan leopard is the island's apex predator, living both within and outside protected areas. In unprotected, shared landscapes, it is important to understand leopard diet and predation patterns to foster long‐term human–leopard coexistence. This study examines the diet of leopards in the human‐dominated tea estate landscape of the Upper Kelani River Basin, in Sri Lanka's Central Highlands. Study goals were to evaluate prey composition, diversity, importance and selection, and to investigate the role of domestic species in leopard diet here. Analysis of 107 leopard scat samples showed leopard feeding behavior was best characterized as generalist and opportunistic, with a wide‐ranging diet ( H ′ = 2.89) consisting of 17 evenly consumed ( D = 0.94) prey species. While the diminutive black‐naped hare (2.5 kg) was most available and most frequently detected in the diet (19.8% of samples), the importance of medium‐sized prey was highlighted, with barking deer (25.5 kg) well utilized (13.9% of samples), representing > 20% of total biomass consumed and showing positive selection (0.281). Moderate selectivity was observed for sambar (0.410), the system's largest potential prey (160–215 kg), which may be expected for meso‐carnivores in the absence of dominant intraguild competition. Primates are a key resource here (23% of samples and biomass) despite being uncommon in tea estates, suggesting preference by leopards. Targeted research to quantify primate abundance and selection is recommended. Overall, wild species represented > 85% of leopard diet, suggesting the landscape retains a substantial natural prey base. Domestic dogs, though common and widely perceived as targeted by leopards here, were moderately avoided (−0.378), a positive outcome for human–leopard coexistence. These findings highlight the leopards' generalist predation tendencies, while suggesting additional complexity and signaling selectivity in predation patterns. Results underscore the necessity of preserving wild prey abundance and diversity to facilitate coexistence in anthropogenically transformed environments.

As humans and wildlife increasingly overlap on a rapidly changing planet, it is important to understand co-existence mechanisms in unprotected landscapes, especially for large carnivores with which humans share an uneasy history. In Sri Lanka’s tea estate-dominated Central Highlands, a dense, widespread human population shares the landscape with the island’s apex predator, the Sri Lankan leopard. This study used remote cameras to determine leopard density, generalized linear models to investigate leopard space-use in relation to key anthropogenic, ecological and environmental variables, and fitted kernel density curves to identify temporal activity patterns in this unprotected, human-dominated region. Adult leopard density (8.82/100 km²) was lower here than in key Sri Lankan National Parks, but not significantly so, suggesting good leopard habitat suitability. Spatially, leopards did not avoid humans but incorporated anthropogenic factors into space-use decisions, with areas of intermediate human and dog presence disproportionately used. Male leopards were more likely to occur at intermediate distances from human settlements, perhaps balancing risk with efficiency while navigating this human-dominated landscape. Female leopards biased space-use towards areas of high natural prey availability, aligning with their need to provide for cubs. Temporally, leopards were significantly less diurnal and more active at dusk than their protected counterparts, likely due to temporal partitioning from diurnally active humans, an adaptation which allows for human-leopard co-existence. This is the first comprehensive study of leopards outside protected areas in Sri Lanka, and highlights the value of unprotected landscapes and their importance for long-term leopard conservation. To foster human-leopard co-existence it is necessary to maintain current human activity patterns which allow for temporal partitioning; ensure existing natural prey resources remain to prevent leopards switching to domestic prey species; and reduce/eliminate the use of wire snares which transcend the effect of temporal partitioning by de-coupling persecution risk from human presence.

Diverse factors, including environmental features and cognitive processes, can drive animals’ movements and space use, with far-reaching implications. For example, repeated use of individual-level travel routeways (directionally constrained but imperfectly aligned routes), which results in spatial concentration of activity, can shape encounter-based processes including predation, mate finding, and disease transmission. However, how much variation in routeway usage exists across species remains unknown. By analyzing GPS movement tracks for 1,239 range-resident mammalian carnivores—representing 16 canid and 18 felid species from six continents—we found strong evidence of a clade-level difference in species’ reliance on repeatedly used travel routeways. Across the global dataset, tracked canids had a 15% (±7 CI) greater density of routeways within their home ranges than did felids, rising to 33% (±16 CI) greater in landscapes shared with tracked felids. Moreover, comparisons within species across landscapes revealed broadly similar home range routeway densities despite habitat differences. On average, canids also reused their travel routeways more intensively than did felids, with hunting strategies and spatial contexts also contributing to the intensity of routeway usage. Collectively, our results suggest that key aspects of carnivore routeway-usage have an evolutionary component. Striking interspecific and clade-level differences in carnivores’ reliance on reused travel routeways within home ranges identify important ways in which the movement patterns of real-world predators depart from classical assumptions of predator-prey theory. Because such departures can drive key aspects of human-wildlife interactions and other encounter-based processes, continued investigations of the relationships between movement mechanisms and space use are critical.

The Sri Lankan leopard (Panthera pardus kotiya) is a threatened endemic sub‐species and the island's apex predator. Wire snares represent a substantial threat to leopards here, with conservation officials routinely rescuing snared leopards. This process can lead to injury treatment, rehabilitation and translocation, the final outcomes of which are poorly understood. Here we report on the rescue and ex situ treatment and recovery of a snared young adult male leopard as well as its post‐recovery release and monitoring. This was the first GPS radio‐collared translocated leopard in Sri Lanka. The leopard was snared in a human‐dominated, unprotected region of the Central Highlands and underwent surgery and 4 months of treatment before being released within a lower elevation protected area ~20 km from the capture site. The leopard made immediate, clear and rapid directional movements towards its original range, island‐hopping by swimming across open water on three occasions (distance range: 90–650 m) to return, within 3 weeks, to a human‐dominated, agricultural landscape ~10 km from its original capture site. Practical Implication: Although the leopard's age and sex, and characteristics of the release suggest a reasonable probability of success, longer monitoring was required to determine the ultimate success of the release. The leopard's immediate attempt to return towards its capture location supports previous research indicating that translocation should be avoided unless circumstances allow for little other choice.

Livestock depredation by wild carnivores threatens carnivore populations and livestock-dependent human communities globally. Understanding local attitudes towards carnivores can inform strategies to improve coexistence. In Sri Lanka, the dairy industry is expanding, creating a need for proactive conflict mitigation. Livestock depredation by the Endangered Sri Lankan leopard Panthera pardus kotiya occurs, but little is known about these incidents or the attitudes of those whose livelihoods may be threatened by this. We surveyed people in two rural communities, Palatupana and Maskeliya, that differed in the scale of livestock ownership, livestock management practices and socio-ecological factors, to characterize attitudes towards leopards and understand their determinants. In Palatupana, an area with extensive cattle rearing, depredation incidents were frequent, and attitudes towards leopards were positively related to respondents' age, number of dependants, years spent rearing livestock and a greater overall support for wildlife conservation. Attitudes were negatively related to respondents' knowledge of leopard ecology and awareness of leopard-related tourism, from which cattle owners do not benefit. In Maskeliya, where cattle rearing is secondary to other agricultural work, depredation did not occur. Here, attitudes were positively related to a desire for increased government assistance with cattle rearing. The inability to develop land for cattle husbandry was a common barrier experienced in both communities. Considering local attitudes can inform strategies to improve human–carnivore coexistence. Approaches that could improve attitudes towards leopards include involvement of cattle owners in tourism programmes, exploring potential alternative land ownership schemes, and improving infrastructure and access to veterinary care.

Predation is a fundamental ecological process influencing the distribution and abundance of animal populations and underlying how prey species perceive risk. The predation process is composed of four sequential stages – search, encounter, attack and kill – each of which has been used to describe risk across the landscape. Here, we used direct observational data of free‐ranging, radio‐collared African lions in Serengeti National Park's western corridor to (1) investigate daily and seasonal predation stage probabilities and (2) using two analytical approaches, compare four mechanisms – prey distribution, intra‐specific competition, spatially anchored landscape features and predator hunting method – that potentially drive spatial predation stage patterns. Results showed that lions encountered potential prey at night significantly less than during diurnal or crepuscular periods. Nocturnal observations were predominantly during full‐moon phases, so if this lower nocturnal encounter rate was due to moon phase it may contribute to lions' typically poor full‐moon hunting success. Predation stage probabilities did not differ between seasons despite high variability in seasonal prey abundance. Spatially, lions encountered potential prey in prey‐rich, open areas near water and spatial range centres. Compared with available areas within seasonal ranges, lion attacks were more likely where prey abundance was high, and kill locations were associated with prey‐rich areas near water and range centres, collectively suggesting opportunistic hunting. However, compared with preceding predation stage locations, attacks occurred near range peripheries and kills where hunting cover was greater, suggesting ambush predation. Our results indicate substantial temporal and spatial variation across the different stages of the predation process. They also highlight first, that results can vary in important ways depending on how analyses are approached, and second, that understanding predator‐prey dynamics depends on analyses of the different stages of predation.

Apex carnivores are integral to effectively functioning ecosystems, but their populations are declining worldwide. To ensure the long‐term viability of top carnivore populations, it is important to understand their ecology and behavior throughout their remaining range, including in protected areas, as these can act as vital refuges. In Sri Lanka, an endangered, endemic leopard sub‐species (Panthera pardus kotiya) is the apex predator, and while its island‐wide distribution is influenced by forest cover and protection level of the landscape, ecological and anthropogenic influences of finer scale landscape utilization are poorly understood. In the interior of Sri Lanka's largest protected area, Wilpattu National Park, we used remote cameras and a spatially explicit capture‐recapture methodology to estimate leopard population density and quantify relative prey availability. We then employed linear models to compare four alternative hypotheses proposed to identify factors influencing observed leopard habitat selection within the PA. Adult leopard density was estimated at 10.4 individuals/100 km²which is at the higher end of the global spectrum of recorded leopard densities, and only marginally lower than other Sri Lankan National Parks which appear to have considerably higher available prey abundance. Leopard site utilization increased with increasing distance from park boundaries, suggesting the presence of edge effects due to anthropogenic disturbance. Leopard occurrence was also higher close to grazing lawns used by prey but not areas of higher prey abundance, evidence of ‘leap‐frogging’, whereby carnivores bias movement toward spatially anchored preferred resources of their prey instead of prey directly. The relatively high core area leopard density, combined with avoidance of boundary areas and frequent observation of injuries suggests the possibility of density‐dependent intra‐specific competition and needs to be further investigated. These results illuminate essential aspects of leopard ecology in a key part of its remaining Sri Lankan range, providing important insights for protected area management.

One of the most challenging tasks in wildlife conservation and management is to clarify how spatial variation in land cover due to anthropogenic disturbance influences wildlife demography and long‐term viability. To evaluate this, we compared rates of survival and population growth by woodland caribou (Rangifer tarandus caribou) from 2 study sites in northern Ontario, Canada that differed in the degree of anthropogenic disturbance because of commercial logging and road development, resulting in differences in predation risk due to gray wolves (Canis lupus). We used an individual‐based model for population viability analysis (PVA) that incorporated adaptive patterns of caribou movement in relation to predation risk and food availability to predict stochastic variation in rates of caribou survival. Field estimates of annual survival rates for adult female caribou in the unlogged ( 0.90) and logged ( 0.76) study sites recorded during 2010–2014 did not differ significantly (P > 0.05) from values predicted by the individual‐based PVA model (unlogged: = 0.87; logged: 0.79). Outcomes from the individual‐based PVA model and a simpler stage‐structured matrix model suggest that substantial differences in adult survival largely due to wolf predation are likely to lead to long‐term decline of woodland caribou in the commercially logged landscape, whereas the unlogged landscape should be considerably more capable of sustaining caribou. Estimates of population growth rates (λ) for the 2010–2014 period differed little between the matrix model and the individual‐based PVA model for the unlogged (matrix model = 1.01; individual‐based model = 0.98) and logged landscape (matrix model = 0.88; individual‐based model = 0.89). We applied the spatially explicit PVA model to assess the viability of woodland caribou across 14 woodland caribou ranges in Ontario. Outcomes of these simulations suggest that woodland caribou ranges that have experienced significant levels of commercial forestry activities in the past had annual growth rates 0.96. These differences were strongly related to regional variation in wolf densities. Our results suggest that increased wolf predation risk due to anthropogenic disturbance is of sufficient magnitude to cause appreciable risk of population decline in woodland caribou in Ontario. © 2020 The Authors. The Journal of Wildlife Management published by Wiley Periodicals, Inc. on behalf of The Wildlife Society.

Context: Differences in body size and mouth morphologies influence dietary resource separation among savanna ungulates, and this influences their distribution across landscape. Aim: The aim was to understand the influence of body size and mouth morphology differences on both diet and patch selection by ungulate species in western Serengeti. Two hypotheses were tested: (1) for ungulate species, the relative body sizes and muzzle widths (mean muzzle width range: 3.1- 9.85) relate directly to food biomass, and to quality of diets selected in the wet season when food is abundant; (2) in the wet season, if food is not limiting, similar-sized species should exhibit greater dietary niche overlap than ungulate species that differ greatly in body size and muzzle width; moreover, similar-sized species exhibit less dietary niche overlap than ungulate species that differ greatly in body size. Methods: In the western Serengeti ecosystem, road transects and direct observation were used to obtain data on the distribution and diet of five ungulate species namely buffalo, zebra, wildebeest, topi and impala; which have of varying sizes (range: 70–630 kg) feeding in three different vegetation types. Grassland biomass, structure, nutrient content and ungulate use were measured at sites along transects. Key Results: Results indicated that large-bodied ungulates utilised patches of greater food abundance compared with those of smaller ungulates. Body mass was also negatively correlated with diet quality, so that smaller animals ate higher protein and lower-fibre foods, as predicted. Diet niche overlap (niche similarity) showed a strong positive relationship with body mass differences among ungulate species, in support of the second of the two predictions from this hypothesis, namely that dissimilarly sized species could eat the same food. Conclusion: Overall, the results suggest that in this savanna system, variation in ungulate body size influences resource separation even in the food-abundant wet season, and that this helps multiple species to co-exist. Implications: Implementing more focused conservation strategies will improve wildlife habitat quality by integrating fire as a forage management tool with grazing preferences to promote forage heterogeneity in protected areas.