Links in a sink: Interplay between habitat structure, ecological constraints and interactions with humans can influence connectivity conservation for tigers in forest corridors.

Global land-use changes and rapid infrastructure development necessitate identification and conservation of wildlife corridors. Connectivity through corridors is shaped by species' structural, ecological and behavioral constraints. In multi-use landscapes, species' interactions with humans could additionally influence connectivity. Using the tiger Panthera tigris as a case study, we make simultaneous assessments of potential connectivity, habitat use and examine their links with the species' negative interactions with humans in central India. We assessed potential connectivity across 10, 000 sq. km of the Kanha-Pench forest corridor using graph-theoretic methods. Combining indirect sign surveys and occupancy models, we examined habitat use, and evaluated its congruence with potential connectivity. Next, we estimated spatial probabilities of livestock depredation through application of multi-state occupancy models to interview-based survey data from local residents. Habitat use by tigers was negatively associated with forest fragmentation and anthropogenic disturbance. Livestock depredation was positively associated with size of settlements and areas most frequented by tigers, and negatively with anthropogenic disturbance within forests. We found high congruence between connectivity and habitat use (r = 0.80); but the strong correlation did not hold in areas with very high levels of livestock depredation levels. Our results indicate that when areas of high use by tigers are constrained by limited connectivity, there are higher chances of human-tiger conflict, and these areas may be ecological traps for the species. Interactions with humans can be crucial in mediating connectivity for large carnivores in shared habitats. Our findings present an opportunity to consolidate areas where carnivore conservation and local livelihood needs can be balanced. Our framework also provides a foundation for spatial prioritization that incorporates a plurality of dimensions, with utility for connectivity conservation of other wide-ranging carnivores.

Human casualties are the dominant cost of human-wildlife conflict in India.

Reducing the costs from human-wildlife conflict, mostly borne by marginal rural households, is a priority for conservation. We estimate the mean species-specific cost for households suffering damages from one of 15 major species of wildlife in India. Our data are from a survey of 5,196 households living near 11 wildlife reserves in India, and self-reported annual costs include crop and livestock losses and human casualties (injuries and death). By employing conservative estimates from the literature on the value of a statistical life (VSL), we find that costs from human casualties overwhelm crop and livestock damages for all species associated with fatalities. Farmers experiencing a negative interaction with an elephant over the last year incur damages on average that are 600 and 900 times those incurred by farmers with negative interactions with the next most costly herbivores: the pig and the nilgai. Similarly, farmers experiencing a negative interaction with a tiger over the last year incur damage that is on average 3 times that inflicted by a leopard and 100 times that from a wolf. These cost differences are largely driven by differences in the incidence of human death and casualties. Our estimate of costs fluctuates across reserves, mostly due to a variation of human casualties. Understanding the drivers of human casualties and reducing their incidence are crucial to reducing the costs from human-wildlife conflict.Most of the tales were about animals, for the Jungle was always at their door. The deer and the pig grubbed up their crops, and now and again the tiger carried off a man at twilight, within sight of the village gates. "Tiger! Tiger!" (Rudyard Kipling, The Jungle Book, Collins Classics, 2010).

Genetic analyses reveal population structure and recent decline in leopards (Panthera pardus fusca) across the Indian subcontinent.

BACKGROUND: Large carnivores maintain the stability and functioning of ecosystems. Currently, many carnivore species face declining population sizes due to natural and anthropogenic pressures. The leopard, Panthera pardus, is probably the most widely distributed and highly adaptable large felid globally, still persisting in most of its historic range. However, we lack subspecies-level data on country or regional scale on population trends, as ecological monitoring approaches are difficult to apply on such wide-ranging species. We used genetic data from leopards sampled across the Indian subcontinent to investigate population structure and patterns of demographic decline. METHODS: We collected faecal samples from the Terai-Arc landscape of northern India and identified 56 unique individuals using a panel of 13 microsatellite markers. We merged this data with already available 143 leopard individuals and assessed genetic structure at country scale. Subsequently, we investigated the demographic history of each identified subpopulations and compared genetic decline analyses with countrywide local extinction probabilities. RESULTS: Our genetic analyses revealed four distinct subpopulations corresponding to Western Ghats, Deccan Plateau-Semi Arid, Shivalik and Terai region of the north Indian landscape, each with high genetic variation. Coalescent simulations with microsatellite loci revealed a possibly human-induced 75-90% population decline between ∼120-200 years ago across India. Population-specific estimates of genetic decline are in concordance with ecological estimates of local extinction probabilities in these subpopulations obtained from occupancy modeling of the historic and current distribution of leopards in India. CONCLUSIONS: Our results confirm the population decline of a widely distributed, adaptable large carnivore. We re-iterate the relevance of indirect genetic methods for such species in conjunction with occupancy assessment and recommend that detailed, landscape-level ecological studies on leopard populations are critical to future conservation efforts. Our approaches and inference are relevant to other widely distributed, seemingly unaffected carnivores such as the leopard.

Extensive vegetation browning and drying in forests of India's Tiger Reserves.

Forest conservation includes stemming deforestation as well as preserving its vegetation condition. Traditional Protected Area (PA) effectiveness evaluations have assessed changes in forest extent but have mostly ignored vegetation condition. Tiger Reserves (TRs) are India's PAs with highest protection and management resources. We used a before-after-control-impact-style design with long-term Landsat 5 TM data to evaluate the effects of protection elevation on vegetation condition (greenness and moisture) in 25 TRs. After declaration as TRs, vegetation condition in 13 TRs (52%) declined in more than 50% of their areas, with 12 TRs (48%) being overall better than their matched Wildlife Sanctuaries (WLSs; PAs with lower protection). In 8 of these TRs analysed for change from before to after declaration, vegetation condition in 5 TRs was harmed over more than 25% of their areas, with 3 TRs being overall better than their matched WLSs. Our results indicate extensive vegetation browning and drying in about half of the study TRs, with these trends often being similar or worse than in matched WLSs. These results suggest that TRs' elevated protection alone may be insufficient to preserve vegetation condition and cast doubt on the effectiveness of protection elevation alone in safeguarding long-term viability of tiger habitats.

Examining human-carnivore interactions using a socio-ecological framework: sympatric wild canids in India as a case study.

Many carnivores inhabit human-dominated landscapes outside protected reserves. Spatially explicit assessments of carnivore distributions and livestock depredation patterns in human-use landscapes are crucial for minimizing negative interactions and fostering coexistence between people and predators. India harbours 23% of the world's carnivore species that share space with 1.3 billion people in approximately 2.3% of the global land area. We examined carnivore distributions and human-carnivore interactions in a multi-use forest landscape in central India. We focused on five sympatric carnivore species: Indian grey wolf Canis lupus pallipes, dhole Cuon alpinus, Indian jackal Canis aureus indicus, Indian fox Vulpes bengalensis and striped hyena Hyaena hyaena. Carnivore occupancy ranged from 12% for dholes to 86% for jackals, mostly influenced by forests, open scrublands and terrain ruggedness. Livestock/poultry depredation probability in the landscape ranged from 21% for dholes to greater than 95% for jackals, influenced by land cover and livestock- or poultry-holding. The five species also showed high spatial overlap with free-ranging dogs, suggesting potential competitive interactions and disease risks, with consequences for human health and safety. Our study provides insights on factors that facilitate and impede co-occurrence between people and predators. Spatial prioritization of carnivore-rich areas and conflict-prone locations could facilitate human-carnivore coexistence in shared habitats. Our framework is ideally suited for making socio-ecological assessments of human-carnivore interactions in other multi-use landscapes and regions, worldwide.

Responses of aerial insectivorous bats to local and landscape-level features of coffee agroforestry systems in Western Ghats, India.

Shade coffee has shown great promise in providing crucial habitats for biodiversity outside formal protected areas. Insectivorous bats have been understudied in coffee, although they may provide pest control services. We investigated the influence of local and landscape-level features of coffee farms on aerial insectivorous bats in Chikmagalur district in the Western Ghats biodiversity hotspot, India. Bats were monitored in 20 farm sites using ultrasound detectors, and the response of bat species richness and activity to changes in tree density, proportion of built-up area in the neighborhood, and distance of farm from forest areas quantified. We examined if models built to explain the species richness and activity could also predict them in nine additional sites. We detected nine phonic types/species in the study area. The quantified predictors had no effect on assemblage-level species richness and activity of bats. Responses of edge-space and cluttered-space forager guilds mirrored those of the overall assemblage, but some species vulnerable to forest conversion like Rhinolophus beddomei were detected rarely. Best models explained up to 20% and 15% variation in assemblage-level species richness and activity respectively, and were poor predictors of both response variables. We conclude that coffee farms in our study area offer an important commuting space for insectivorous bats across a gradient of shade management. Further research should include species-specific responses to management decisions for at-risk species and quantification of ecosystem services like natural pest control to inform biodiversity conservation initiatives in the Western Ghats coffee landscapes.

Author Correction: Birds and beans: Comparing avian richness and endemism in arabica and robusta agroforests in India's Western Ghats.

A correction to this article has been published and is linked from the HTML and PDF versions of this paper. The error has not been fixed in the paper.

Birds and beans: Comparing avian richness and endemism in arabica and robusta agroforests in India's Western Ghats.

Coffee is a major tropical commodity crop that can provide supplementary habitat for native wildlife. In Asia, coffee production is an increasingly important driver of landscape transformation and shifts between different coffee species is a major dimension of agroforestry trends. Yet few studies have compared the ecological impacts of conversion between different coffee species. We evaluated whether or not the two species of coffee grown globally-Coffea arabica and C. canephora (denoted "robusta")-had equivalent avian conservation value in the Western Ghats, India, where robusta production has become increasingly dominant. We found that habitat specialist and functional guild diversity was higher in arabica, and that arabica was more profitable. However, robusta farms generally supported the same or slightly higher abundances of habitat specialists and functional guilds, largely due to dense canopy and landscape-level forest cover. Farming practices, chiefly pesticide use, may affect the suitability of coffee agroforests as habitat for avian specialists, and at present, robusta farmers tended to use less pesticide. Given future projections for arabica to robusta conversion in tropical Asia, our study indicates that certification efforts should prioritize maintaining native canopy shade trees and forest cover to ensure that coffee landscapes can continue providing biodiversity benefits.

Wildlife in the Matrix: Spatio-Temporal Patterns of Herbivore Occurrence in Karnataka, India.

Wildlife reserves are becoming increasingly isolated from the surrounding human-dominated landscapes particularly in Asia. It is imperative to understand how species are distributed spatially and temporally in and outside reserves, and what factors influence their occurrence. This study surveyed 7500 km(2) landscape surrounding five reserves in the Western Ghats to examine patterns of occurrence of five herbivores: elephant, gaur, sambar, chital, and pig. Species distributions are modeled spatio-temporally using an occupancy approach. Trained field teams conducted 3860 interview-based occupancy surveys in a 10-km buffer surrounding these five reserves in 2012. I found gaur and wild pig to be the least and most wide-ranging species, respectively. Elephant and chital exhibit seasonal differences in spatial distribution unlike the other three species. As predicted, distance to reserve, the reserve itself, and forest cover were associated with higher occupancy of all species, and higher densities of people negatively influenced occurrence of all species. Park management, species protection, and conflict mitigation efforts in this landscape need to incorporate temporal and spatial understanding of species distributions. All species are known crop raiders and conflict prone locations with resources (such as water and forage) have to be monitored and managed carefully. Wildlife reserves and adjacent areas are critical for long-term persistence and habitat use for all five herbivores and must be monitored to ensure wildlife can move freely. Such a large-scale approach to map and monitor species distributions can be adapted to other landscapes to identify and monitor critical habitats shared by people and wildlife.

Spotted in the News: Using Media Reports to Examine Leopard Distribution, Depredation, and Management Practices outside Protected Areas in Southern India.

There is increasing evidence of large carnivore presence outside protected areas, globally. Although this spells conservation success through population recoveries, it makes carnivore persistence in human-use landscapes tenuous. The widespread distribution of leopards in certain regions of India typifies this problem. We obtained information on leopard-human interactions at a regional scale in Karnataka State, India, based on systematic surveys of local media reports. We applied an innovative occupancy modelling approach to map their distribution patterns and identify hotspots of livestock/human depredation. We also evaluated management responses like removals of 'problem' leopards through capture and translocations. Leopards occupied around 84,000 km2 or 47% of the State's geographic area, outside designated national parks and wildlife sanctuaries. Their presence was facilitated by extent of vegetative cover- including irrigated croplands, rocky escarpments, and prey base in the form of feral and free-ranging dogs. Higher probabilities of livestock/human attacks by leopards were associated with similar ecological features as well as with capture/removals of leopards. Of the 56 cases of leopard removals reported, 91% did not involve human attacks, but followed livestock predation or only leopard sightings. The lack of knowledge on leopard ecology in human-use areas has resulted in unscientific interventions, which could aggravate the problem rather than mitigating it. Our results establish the presence of resident, breeding leopards in human-use areas. We therefore propose a shift in management focus, from current reactive practices like removal and translocation of leopards, to proactive measures that ensure safety of human lives and livelihoods.

Patterns and Determinants of Habitat Occupancy by the Asian Elephant in the Western Ghats of Karnataka, India.

Understanding species distribution patterns has direct ramifications for the conservation of endangered species, such as the Asian elephant Elephas maximus. However, reliable assessment of elephant distribution is handicapped by factors such as the large spatial scales of field studies, survey expertise required, the paucity of analytical approaches that explicitly account for confounding observation processes such as imperfect and variable detectability, unequal sampling probability and spatial dependence among animal detections. We addressed these problems by carrying out 'detection--non-detection' surveys of elephant signs across a c. 38,000-km(2) landscape in the Western Ghats of Karnataka, India. We analyzed the resulting sign encounter data using a recently developed modeling approach that explicitly addresses variable detectability across space and spatially dependent non-closure of occupancy, across sampling replicates. We estimated overall occupancy, a parameter useful to monitoring elephant populations, and examined key ecological and anthropogenic drivers of elephant presence. Our results showed elephants occupied 13,483 km(2) (SE = 847 km(2)) corresponding to 64% of the available 21,167 km(2) of elephant habitat in the study landscape, a useful baseline to monitor future changes. Replicate-level detection probability ranged between 0.56 and 0.88, and ignoring it would have underestimated elephant distribution by 2116 km(2) or 16%. We found that anthropogenic factors predominated over natural habitat attributes in determining elephant occupancy, underscoring the conservation need to regulate them. Human disturbances affected elephant habitat occupancy as well as site-level detectability. Rainfall is not an important limiting factor in this relatively humid bioclimate. Finally, we discuss cost-effective monitoring of Asian elephant populations and the specific spatial scales at which different population parameters can be estimated. We emphasize the need to model the observation and sampling processes that often obscure the ecological process of interest, in this case relationship between elephants to their habitat.

On a dhole trail: examining ecological and anthropogenic correlates of dhole habitat occupancy in the Western ghats of India.

Although they play a critical role in shaping ecological communities, many threatened predator species are data-deficient. The Dhole Cuon alpinus is one such rare canid with a global population thought to be <2500 wild individuals. We assessed habitat occupancy patterns of dholes in the Western Ghats of Karnataka, India, to understand ecological and anthropogenic determinants of their distribution and habitat-use. We conducted spatially replicated detection/non-detection surveys of dhole signs along forest trails at two appropriate scales: the entire landscape and a single wildlife reserve. Landscape-scale habitat occupancy was assessed across 38,728 km(2) surveying 206 grid cells of 188-km(2) each. Finer scale habitat-use within 935 km2 Bandipur Reserve was studied surveying 92 grid cells of 13-km(2) km each. We analyzed the resulting data of dhole signs using likelihood-based habitat occupancy models. The models explicitly addressed the problematic issue of imperfect detection of dhole signs during field surveys as well as potential spatial auto-correlation between sign detections made on adjacent trail segments. We show that traditional 'presence versus absence' analyses underestimated dhole habitat occupancy by 60% or 8682 km2 [naïve = 0.27; ψL(SE)  = 0.68 (0.08)] in the landscape. Addressing imperfect sign detections by estimating detection probabilities [p(t)(L) (SE) = 0.12 (0.11)] was critical for reliable estimation. Similar underestimation occurred while estimating habitat-use probability at reserve-scale [naïve = 0.39; Ψs(SE) = 0.71 (0.06)]. At landscape scale, relative abundance of principal ungulate prey primarily influenced dhole habitat occupancy. Habitat-use within a reserve, however, was predominantly and negatively influenced by anthropogenic disturbance. Our results are the first rigorous assessment of dhole occupancy at multiple spatial scales with potential conservation value. The approach used in this study has potential utility for cost-effectively assessing spatial distribution and habitat-use in other species, landscapes and reserves.

Living with wildlife and mitigating conflicts around three Indian protected areas.

Crop and livestock losses to wildlife are a concern for people neighboring many protected areas (PAs) and can generate opposition to conservation. Examining patterns of conflict and associated tolerance is important to devise policies to reduce conflict impacts on people and wildlife. We surveyed 398 households from 178 villages within 10 km of Ranthambore, Kanha, and Nagarahole parks in India. We compared different attitudes toward wildlife, and presented hypothetical response scenarios, including killing the problem animal(s). Eighty percent of households reported crop losses to wildlife and 13 % livestock losses. Higher crop loss was associated with more cropping months per year, greater crop variety, and more harvest seasons per year but did not vary with proximity to the PA, suggesting that PAs are not necessarily "sources" for crop raiders. By contrast, complaints of "depredating carnivores" were associated with people-grazing animals and collecting resources from PAs. Many households (83 %) engaged in mitigation efforts. We found that only fencing and guard animals reduce crop losses, and no efforts to lower livestock losses. Contrary to our expectations, carnivores were not viewed with more hostility than crop-raiding wildlife. Households reported greater inclination to kill herbivores destroying crops or carnivores harming people, but not carnivores preying on livestock.Our model estimated probability of [corrected] crop loss was 82 % across surveyed households (highest in Kanha),while the livestock loss experienced was 27 % (highest in Ranthambore). Our comparative study provides insights into factors associated with conflict loss and tolerance, and aids in improving ongoing conservation and compensation efforts.

Assessing patterns of human-wildlife conflicts and compensation around a Central Indian protected area.

Mitigating crop and livestock loss to wildlife and improving compensation distribution are important for conservation efforts in landscapes where people and wildlife co-occur outside protected areas. The lack of rigorously collected spatial data poses a challenge to management efforts to minimize loss and mitigate conflicts. We surveyed 735 households from 347 villages in a 5154 km(2) area surrounding Kanha Tiger Reserve in India. We modeled self-reported household crop and livestock loss as a function of agricultural, demographic and environmental factors, and mitigation measures. We also modeled self-reported compensation received by households as a function of demographic factors, conflict type, reporting to authorities, and wildlife species involved. Seventy-three percent of households reported crop loss and 33% livestock loss in the previous year, but less than 8% reported human injury or death. Crop loss was associated with greater number of cropping months per year and proximity to the park. Livestock loss was associated with grazing animals inside the park and proximity to the park. Among mitigation measures only use of protective physical structures were associated with reduced livestock loss. Compensation distribution was more likely for tiger related incidents, and households reporting loss and located in the buffer. Average estimated probability of crop loss was 0.93 and livestock loss was 0.60 for surveyed households. Estimated crop and livestock loss and compensation distribution were higher for households located inside the buffer. Our approach modeled conflict data to aid managers in identifying potential conflict hotspots, influential factors, and spatially maps risk probability of crop and livestock loss. This approach could help focus allocation of conservation efforts and funds directed at conflict prevention and mitigation where high densities of people and wildlife co-occur.

Local residents perception of benefits and losses from protected areas in India and Nepal.

High densities of people living around protected areas (PAs) in South Asia require management strategies to balance conservation goals and livelihood needs. Based on a survey of 777 households around five PAs in India and Nepal, this paper provides a comparative perspective of Indian and Nepali households' views of protected area benefits and costs, their attitude toward conservation in general, and attitude toward protected area staff. Results indicate mixed responses towards tourism, varying from very favorable in Nepal to less favorable in India. The majority (81%) held positive attitudes towards the existence and importance of PAs but had negative perceptions of PA staff (69%). Most residents perceived benefits from access to fuel wood, fodder and other PA resources including benefits from tourism, while crop and livestock losses from wildlife were the main costs. Households overall positive attitudes towards the PAs and conservation despite high losses from living around PAs suggests that local residents may support conservation if their livelihood needs are met. Comparisons of household attitudes and perceptions suggest that locally based strategies rather than top-down approaches are likely to be more effective. Extending PA benefits to smaller landholders, households that are highly resource-dependent or experiencing higher income losses from human-wildlife conflicts, and less educated residents are particularly important to balance costs and losses from living around protected areas.

The shrinking ark: patterns of large mammal extinctions in India.

Mammal extinctions are widespread globally, with South Asian species being most threatened. We examine local extinctions of 25 mammals in India. We use historical records to obtain a set of locations at which each species was known to have been present at some time in the last 200 years. We then use occupancy estimation models to draw inferences about current presence at these same locations based on field observations of local experts. We examine predictions about the influence of key factors such as protected areas, forest cover, elevation, human population density and cultural tolerance on species extinction. For all 25 species, estimated local extinction probabilities (referenced to a 100 year time frame) range between 0.14 and 0.96. Time elapsed since the historical occurrence record was an important determinant of extinction probability for 14 species. Protected areas are positively associated with lower extinction of 18 species, although many species occur outside them. We find evidence that higher proportion of forest cover is associated with lower extinction probabilities for seven species. However, for species that prefer open habitats (which have experienced intensive land-use change), forest cover alone appears insufficient to ensure persistence (the complement of extinction). We find that higher altitude is positively associated with lower extinction for eight species. Human population density is positively associated with extinction of 13 species. We find that 'culturally tolerated' species do exhibit higher persistence. Overall, large-bodied, rare and habitat specialist mammals tend to have higher extinction probabilities.