Topcats and underdogs: intraguild interactions among three apex carnivores across Asia's forestscapes.

Intraguild interactions among carnivores have long held the fascination of ecologists. Ranging from competition to facilitation and coexistence, these interactions and their complex interplay influence everything from species persistence to ecosystem functioning. Yet, the patterns and pathways of such interactions are far from understood in tropical forest systems, particularly across countries in the Global South. Here, we examined the determinants and consequences of competitive interactions between dholes Cuon alpinus and the two large felids (leopards Panthera pardus and tigers Panthera tigris) with which they most commonly co-occur across Asia. Using a combination of traditional and novel data sources (N = 118), we integrate information from spatial, temporal, and dietary niche dimensions. These three species have faced catastrophic declines in their extent of co-occurrence over the past century; most of their source populations are now confined to Protected Areas. Analysis of dyadic interactions between species pairs showed a clear social hierarchy. Tigers were dominant over dholes, although pack strength in dholes helped ameliorate some of these effects; leopards were subordinate to dholes. Population-level spatio-temporal interactions assessed at 25 locations across Asia did not show a clear pattern of overlap or avoidance between species pairs. Diet-profile assessments indicated that wild ungulate biomass consumption by tigers was highest, while leopards consumed more primate and livestock prey as compared to their co-predators. In terms of prey offtake (ratio of wild prey biomass consumed to biomass available), the three species together harvested 0.4-30.2% of available prey, with the highest offtake recorded from the location where the carnivores reach very high densities. When re-examined in the context of prey availability and offtake, locations with low wild prey availability showed spatial avoidance and temporal overlap among the carnivore pairs, and locations with high wild prey availability showed spatial overlap and temporal segregation. Based on these observations, we make predictions for 40 Protected Areas in India where temporally synchronous estimates of predator and prey densities are available. We expect that low prey availability will lead to higher competition, and in extreme cases, to the complete exclusion of one or more species. In Protected Areas with high prey availability, we expect intraguild coexistence and conspecific competition among carnivores, with spill-over to forest-edge habitats and subsequent prey-switching to livestock. We stress that dhole-leopard-tiger co-occurrence across their range is facilitated through an intricate yet fragile balance between prey availability, and intraguild and conspecific competition. Data gaps and limitations notwithstanding, our study shows how insights from fundamental ecology can be of immense utility for applied aspects like large predator conservation and management of human-carnivore interactions. Our findings also highlight potential avenues for future research on tropical carnivores that can broaden current understanding of intraguild competition in forest systems of Asia and beyond.

Methodological approaches for estimating populations of the endangered dhole Cuon alpinus.

Large carnivores are important for maintaining ecosystem integrity and attract much research and conservation interest. For most carnivore species, estimating population density or abundance is challenging because they do not have unique markings for individual identification. This hinders status assessments for many threatened species, and calls for testing new methodological approaches. We examined past efforts to assess the population status of the endangered dhole (Cuon alpinus), and explored the application of a suite of recently developed models for estimating their populations using camera-trap data from India's Western Ghats. We compared the performance of Site-Based Abundance (SBA), Space-to-Event (STE), and Time-to-Event (TTE) models against current knowledge of their population size in the area. We also applied two of these models (TTE and STE) to the co-occurring leopard (Panthera pardus), for which density estimates were available from Spatially Explicit Capture-Recapture (SECR) models, so as to simultaneously validate the accuracy of estimates for one marked and one unmarked species. Our review of literature (n = 38) showed that most assessments of dhole populations involved crude indices (relative abundance index; RAI) or estimates of occupancy and area of suitable habitat; very few studies attempted to estimate populations. Based on empirical data from our field surveys, the TTE and SBA models overestimated dhole population size beyond ecologically plausible limits, but the STE model produced reliable estimates for both the species. Our findings suggest that it is difficult to estimate population sizes of unmarked species when model assumptions are not fully met and data are sparse, which are commonplace for most ecological surveys in the tropics. Based on our assessment, we propose that practitioners who have access to photo-encounter data on dholes across Asia test old and new analytical approaches to increase the overall knowledge-base on the species, and contribute towards conservation monitoring of this endangered carnivore.

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.

Multiple coping strategies maintain stability of a small mammal population in a resource-restricted environment.

In semi-arid environments, aperiodic rainfall pulses determine plant production and resource availability for higher trophic levels, creating strong bottom-up regulation. The influence of climatic factors on population vital rates often shapes the dynamics of small mammal populations in such resource-restricted environments. Using a 21-year biannual capture-recapture dataset (1993 to 2014), we examined the impacts of climatic factors on the population dynamics of the brush mouse (Peromyscus boylii) in semi-arid oak woodland of coastal-central California. We applied Pradel's temporal symmetry model to estimate capture probability (p), apparent survival (φ), recruitment (f), and realized population growth rate (λ) of the brush mouse and examined the effects of temperature, rainfall, and El Niño on these demographic parameters. The population was stable during the study period with a monthly realized population growth rate of 0.993 ± SE 0.032, but growth varied over time from 0.680 ± 0.054 to 1.450 ± 0.083. Monthly survival estimates averaged 0.789 ± 0.005 and monthly recruitment estimates averaged 0.175 ± 0.038. Survival probability and realized population growth rate were positively correlated with rainfall and negatively correlated with temperature. In contrast, recruitment was negatively correlated with rainfall and positively correlated with temperature. Brush mice maintained their population through multiple coping strategies, with high recruitment during warmer and drier periods and higher survival during cooler and wetter conditions. Although climatic change in coastal-central California will likely favor recruitment over survival, varying strategies may serve as a mechanism by which brush mice maintain resilience in the face of climate change. Our results indicate that rainfall and temperature are both important drivers of brush mouse population dynamics and will play a significant role in predicting the future viability of brush mice under a changing climate.

Every dog has its prey: Range-wide assessment of links between diet patterns, livestock depredation and human interactions for an endangered carnivore.

Livestock depredation is the most ubiquitous type of negative interaction between humans and carnivores. We conducted a range-wide assessment linking diet patterns of the endangered dhole Cuon alpinus, with livestock consumption and human-dhole interactions. We first performed a reanalysis of dhole diet data from all published studies (1973-2013) incorporating a recently-developed non-linear correction factor for quantifying prey biomass consumed. We then determined the relative livestock numbers consumed by dholes over time across its range, compared these with earlier estimates, and investigated the relative importance of wild vs. non-wild prey in dhole diet. Using information from >70 studies, we explored links between livestock consumption by dholes, availability of wild versus non-wild prey, sympatric depredation-prone carnivores, and people's perception of dholes as livestock predators. We found that (a) dhole diet profiles varied regionally, (b) dholes consumed fewer livestock compared to estimates generated using other, widely used methods, (c) livestock consumption by dholes was associated with wild and non-wild prey densities, and number of co-predator species, and (d) people's negative perception of dholes was associated with pack sizes, levels of livestock depredation and number of sympatric carnivore species. Global efforts for dhole conservation should involve different strategies based on region-specific realities that account for ecological context as well as human perceptions, which would require well-designed studies of dhole social and population dynamics, and human-dhole interactions. We also call for more such range-wide assessments of livestock depredation by wild canids, complemented with direct investigations of human-canid interactions.

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.

Insights from distribution dynamics inform strategies to conserve a dhole Cuon alpinus metapopulation in India.

Most large carnivore populations currently occur in heterogeneous landscapes, with source populations embedded in a matrix of human-dominated habitats. Understanding changes in distribution of endangered carnivores is critical for prioritizing and implementing conservation strategies. We examined distribution and dynamics of a dhole Cuon alpinus metapopulation, first in 2007 and subsequently in 2015, based on indirect sign surveys across 37, 000sq. km of India's Western Ghats. Predicted dhole occupancy declined from 0.62 (95% CI: 0.58-0.66) in 2007 to 0.54 (95% CI: 0.50-0.58) in 2015. Occupancy was associated with abundance of primary prey species and anthropogenic disturbance. Local extinction appeared to be influenced by forest cover loss, and offset by protected reserves; colonization was influenced by occupancy in neighbouring sites. Perturbation analysis indicated that occupancy was more sensitive to local extinction within reserves and to colonization in sites abutting reserves. The Western Ghats could serve as a stronghold for the endangered dhole, provided future colonizations are facilitated through habitat consolidation beyond reserve boundaries, and local extinctions are prevented by increasing protection efforts within select reserves. We advocate for wildlife managers to adopt a landscape-based approach and periodic monitoring to ensure persistence of the dhole metapopulation in Western Ghats, and in other critical conservation regions across the species' geographic range.

Large anthropogenic impacts on a charismatic small carnivore: Insights from distribution surveys of red panda Ailurus fulgens in Nepal.

Protected areas are key to preserving biodiversity and maintaining ecosystem services. However, their ability to ensure long-term survival of threatened andendangered species varies across countries, regions and landscapes. Distribution surveys can beparticularly important for assessing the value of protected areas, and gauging their efficacy incatering to species-specific requirements. We assessed the conservation value of one such reserve for a charismatic yet globally endangered species, the red panda Ailurus fulgens,in the light of on-going land-use transformation in Nepal. We conducted field surveys forindirect signs of red pandas along forest trails in 25-km2 sampling grid cells (n = 54) of Dhorpatan Hunting Reserve, and confronted a set of ecological hypotheses to the data using hierarchical occupancy models. We estimated overall occupancy at Ψ(SE) = 0.41 (0.007), with relatively high site-level detectability [p = 0.93 (SE = 0.001)]. Our results show that despitebeing a subsistence form of small-scale resource use, extraction of bamboo and livestock grazing negatively affected panda occurrence, albeit at different intensities. The amount of bamboo cover,rather than the overall proportion of forest cover, had greater influence on the panda occurrence. Despite availability of bamboo cover, areas with bamboo extraction and anthropogenic disturbances were less likely to be occupied by pandas. Together, these results suggest that long-term persistence of red pandas in this reserve and elsewhere across the species' range will require preventing commercial extractionof bamboo, coupled with case-specific regulation of anthropogenic exploitation of red panda habitats.

Spatio-temporal interactions facilitate large carnivore sympatry across a resource gradient.

Species within a guild vary their use of time, space and resources, thereby enabling sympatry. As intra-guild competition intensifies, such behavioural adaptations may become prominent. We assessed mechanisms of facilitating sympatry among dhole (Cuon alpinus), leopard (Panthera pardus) and tiger (Panthera tigris) in tropical forests of India using camera-trap surveys. We examined population-level temporal, spatial and spatio-temporal segregation among them across four reserves representing a gradient of carnivore and prey densities. Temporal and spatial overlaps were higher at lower prey densities. Combined spatio-temporal overlap was minimal, possibly due to chance. We found fine-scale avoidance behaviours at one high-density reserve. Our results suggest that: (i) patterns of spatial, temporal and spatio-temporal segregation in sympatric carnivores do not necessarily mirror each other; (ii) carnivores are likely to adopt temporal, spatial, and spatio-temporal segregation as alternative mechanisms to facilitate sympatry; and (iii) carnivores show adaptability across a gradient of resource availability, a driver of inter-species competition. We discuss behavioural mechanisms that permit carnivores to co-occupy rather than dominate functional niches, and adaptations to varying intensities of competition that are likely to shape structure and dynamics of carnivore guilds.

Dynamics of a low-density tiger population in Southeast Asia in the context of improved law enforcement.

Recovering small populations of threatened species is an important global conservation strategy. Monitoring the anticipated recovery, however, often relies on uncertain abundance indices rather than on rigorous demographic estimates. To counter the severe threat from poaching of wild tigers (Panthera tigris), the Government of Thailand established an intensive patrolling system in 2005 to protect and recover its largest source population in Huai Kha Khaeng Wildlife Sanctuary. Concurrently, we assessed the dynamics of this tiger population over the next 8 years with rigorous photographic capture-recapture methods. From 2006 to 2012, we sampled across 624-1026 km(2) with 137-200 camera traps. Cameras deployed for 21,359 trap days yielded photographic records of 90 distinct individuals. We used closed model Bayesian spatial capture-recapture methods to estimate tiger abundances annually. Abundance estimates were integrated with likelihood-based open model analyses to estimate rates of annual and overall rates of survival, recruitment, and changes in abundance. Estimates of demographic parameters fluctuated widely: annual density ranged from 1.25 to 2.01 tigers/100 km(2) , abundance from 35 to 58 tigers, survival from 79.6% to 95.5%, and annual recruitment from 0 to 25 tigers. The number of distinct individuals photographed demonstrates the value of photographic capture-recapture methods for assessments of population dynamics in rare and elusive species that are identifiable from natural markings. Possibly because of poaching pressure, overall tiger densities at Huai Kha Khaeng were 82-90% lower than in ecologically comparable sites in India. However, intensified patrolling after 2006 appeared to reduce poaching and was correlated with marginal improvement in tiger survival and recruitment. Our results suggest that population recovery of low-density tiger populations may be slower than anticipated by current global strategies aimed at doubling the number of wild tigers in a decade.

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.

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.