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.

Density estimation in tiger populations: combining information for strong inference.

A productive way forward in studies of animal populations is to efficiently make use of all the information available, either as raw data or as published sources, on critical parameters of interest. In this study, we demonstrate two approaches to the use of multiple sources of information on a parameter of fundamental interest to ecologists: animal density. The first approach produces estimates simultaneously from two different sources of data. The second approach was developed for situations in which initial data collection and analysis are followed up by subsequent data collection and prior knowledge is updated with new data using a stepwise process. Both approaches are used to estimate density of a rare and elusive predator, the tiger, by combining photographic and fecal DNA spatial capture-recapture data. The model, which combined information, provided the most precise estimate of density (8.5 +/- 1.95 tigers/100 km2 [posterior mean +/- SD]) relative to a model that utilized only one data source (photographic, 12.02 +/- 3.02 tigers/100 km2 and fecal DNA, 6.65 +/- 2.37 tigers/100 km2). Our study demonstrates that, by accounting for multiple sources of available information, estimates of animal density can be significantly improved.

Why the Indian subcontinent holds the key to global tiger recovery.

With only approximately 3,000 wild individuals surviving restricted to just 7% of their historical range, tigers are now a globally threatened species. Therefore, conservation efforts must prioritize regions that harbor more tigers, as well try to capture most of the remaining genetic variation and habitat diversity. Only such prioritization based on demographic, genetic, and ecological considerations can ensure species recovery and retention of evolutionary flexibility in the face of ongoing global changes. Although scientific understanding of ecological and demographic aspects of extant wild tiger populations has improved recently, little is known about their genetic composition and variability. We sampled 73 individual tigers from 28 reserves spread across a diversity of habitats in the Indian subcontinent to obtain 1,263 bp of mitochondrial DNA and 10 microsatellite loci. Our analyses reveals that Indian tigers retain more than half of the extant genetic diversity in the species. Coalescent simulations attribute this high genetic diversity to a historically large population size of about 58,200 tigers for peninsular India south of the Gangetic plains. Furthermore, our analyses indicate a precipitous, possibly human-induced population crash approximately 200 years ago in India, which is in concordance with historical records. Our results suggest that only 1.7% (with an upper limit of 13% and a lower limit of 0.2%) of tiger numbers in historical times remain now. In the global conservation context our results suggest that, based on genetic, demographic, and ecological considerations, the Indian subcontinent holds the key to global survival and recovery of wild tigers.

Translocation as a tool for mitigating conflict with leopards in human-dominated landscapes of India.

We examined the efficacy of a translocation program in which large numbers of leopards (Panthera pardus fusca) were trapped in human-dominated landscapes where livestock attacks were common and human attacks rare and released into adjoining forested areas in an attempt to reduce leopard presence and mitigate conflicts at the capture site. In the year starting in February of 2001, 29 leopards were captured in the human-dominated rural landscape of the Junnar region (4275 km(2) , 185 people/km(2) ), Maharashtra, India, and released an average of 39.5 km away in adjoining forests. Eleven leopards were also relocated to the same forests from other districts. Prior to the large-scale translocation program, an average of four leopard attacks on humans occurred each year between 1993 and 2001. After the translocation program was initiated, the average increased substantially to 17 attacks. Linear and logistic models showed that attack frequency increased in Junnar following nearby releases of leopards and decreased when leopards were removed for releases far away; that attacks became more lethal when the number of leopards introduced from other districts increased; and that attacks were most likely to occur in the regions where the largest number of leopards had been introduced from other areas. These results suggest that leopards did not stay at the release sites and that translocation induced attacks on people. Potential explanations for these results include increased aggression induced by stress of the translocation process, movement through unfamiliar human-dominated landscapes following release, and loss of fear of humans due to familiarity with humans acquired during captivity. Our results show that reactive solutions to attacks on humans by leopards, such as translocation, could in fact increase human-leopard conflict. Measures to reduce human-carnivore conflicts may include more effective compensation procedures to pay livestock owners for the loss of animals to predation by carnivores, providing better methods of protection for livestock, and encouraging greater social acceptance of the presence of carnivores in human-dominated landscapes.

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.

Bayesian inference in camera trapping studies for a class of spatial capture-recapture models.

We develop a class of models for inference about abundance or density using spatial capture-recapture data from studies based on camera trapping and related methods. The model is a hierarchical model composed of two components: a point process model describing the distribution of individuals in space (or their home range centers) and a model describing the observation of individuals in traps. We suppose that trap- and individual-specific capture probabilities are a function of distance between individual home range centers and trap locations. We show that the models can be regarded as generalized linear mixed models, where the individual home range centers are random effects. We adopt a Bayesian framework for inference under these models using a formulation based on data augmentation. We apply the models to camera trapping data on tigers from the Nagarahole Reserve, India, collected over 48 nights in 2006. For this study, 120 camera locations were used, but cameras were only operational at 30 locations during any given sample occasion. Movement of traps is common in many camera-trapping studies and represents an important feature of the observation model that we address explicitly in our application.

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.

State space and movement specification in open population spatial capture-recapture models.

With continued global changes, such as climate change, biodiversity loss, and habitat fragmentation, the need for assessment of long-term population dynamics and population monitoring of threatened species is growing. One powerful way to estimate population size and dynamics is through capture-recapture methods. Spatial capture (SCR) models for open populations make efficient use of capture-recapture data, while being robust to design changes. Relatively few studies have implemented open SCR models, and to date, very few have explored potential issues in defining these models. We develop a series of simulation studies to examine the effects of the state-space definition and between-primary-period movement models on demographic parameter estimation. We demonstrate the implications on a 10-year camera-trap study of tigers in India. The results of our simulation study show that movement biases survival estimates in open SCR models when little is known about between-primary-period movements of animals. The size of the state-space delineation can also bias the estimates of survival in certain cases.We found that both the state-space definition and the between-primary-period movement specification affected survival estimates in the analysis of the tiger dataset (posterior mean estimates of survival ranged from 0.71 to 0.89). In general, we suggest that open SCR models can provide an efficient and flexible framework for long-term monitoring of populations; however, in many cases, realistic modeling of between-primary-period movements is crucial for unbiased estimates of survival and density.

Estimating densities of large herbivores in tropical forests: Rigorous evaluation of a dung-based method.

When sighting-based surveys to estimate population densities of large herbivores in tropical dense forests are not practical or affordable, surveys that rely on animal dung are sometimes used. This study tested one such dung-based method by deriving population densities from observed dung densities of six large herbivores (chital, elephant, gaur, muntjac, sambar, and wild pig) in two habitats, dry deciduous forests (DDF) and moist deciduous forests (MDF), within Nagarahole National Park, southern India. Using the program DUNGSURV, dung pile counts, decay rates estimated from field experiments, and defecation rates derived from literature were analyzed together by a model that allows for random events affecting dung decay. Densities of chital were the highest, followed by sambar. Wild pig densities were similar in the two habitats, sambar densities were higher in DDF, and densities of the other species were higher in MDF than in DDF. We compared DUNGSURV estimates with densities estimated using distance sampling in the same season. DUNGSURV estimates were substantially higher for all species in both habitats. These differences highlight the challenges that researchers face in computing unbiased estimates of dung decay rates and in relying on defecation rates from literature. Besides the elephant, this study is the first to rigorously test the efficacy of using a dung-based approach to estimate densities of large herbivore species in Asia, and based on this evaluation, we provide specific recommendations to address issues that require careful consideration before observed dung densities are used to derive animal densities. Our results underline the need for an experimental study of a known population in a fenced reserve to validate the true potential of using dung-based approaches to estimate population densities.

A hierarchical model for estimating the spatial distribution and abundance of animals detected by continuous-time recorders.

MOTIVATION: Several spatial capture-recapture (SCR) models have been developed to estimate animal abundance by analyzing the detections of individuals in a spatial array of traps. Most of these models do not use the actual dates and times of detection, even though this information is readily available when using continuous-time recorders, such as microphones or motion-activated cameras. Instead most SCR models either partition the period of trap operation into a set of subjectively chosen discrete intervals and ignore multiple detections of the same individual within each interval, or they simply use the frequency of detections during the period of trap operation and ignore the observed times of detection. Both practices make inefficient use of potentially important information in the data. MODEL AND DATA ANALYSIS: We developed a hierarchical SCR model to estimate the spatial distribution and abundance of animals detected with continuous-time recorders. Our model includes two kinds of point processes: a spatial process to specify the distribution of latent activity centers of individuals within the region of sampling and a temporal process to specify temporal patterns in the detections of individuals. We illustrated this SCR model by analyzing spatial and temporal patterns evident in the camera-trap detections of tigers living in and around the Nagarahole Tiger Reserve in India. We also conducted a simulation study to examine the performance of our model when analyzing data sets of greater complexity than the tiger data. BENEFITS: Our approach provides three important benefits: First, it exploits all of the information in SCR data obtained using continuous-time recorders. Second, it is sufficiently versatile to allow the effects of both space use and behavior of animals to be specified as functions of covariates that vary over space and time. Third, it allows both the spatial distribution and abundance of individuals to be estimated, effectively providing a species distribution model, even in cases where spatial covariates of abundance are unknown or unavailable. We illustrated these benefits in the analysis of our data, which allowed us to quantify differences between nocturnal and diurnal activities of tigers and to estimate their spatial distribution and abundance across the study area. Our continuous-time SCR model allows an analyst to specify many of the ecological processes thought to be involved in the distribution, movement, and behavior of animals detected in a spatial trapping array of continuous-time recorders. We plan to extend this model to estimate the population dynamics of animals detected during multiple years of SCR surveys.

Assessing tiger population dynamics using photographic capture-recapture sampling.

Although wide-ranging, elusive, large carnivore species, such as the tiger, are of scientific and conservation interest, rigorous inferences about their population dynamics are scarce because of methodological problems of sampling populations at the required spatial and temporal scales. We report the application of a rigorous, noninvasive method for assessing tiger population dynamics to test model-based predictions about population viability. We obtained photographic capture histories for 74 individual tigers during a nine-year study involving 5725 trap-nights of effort. These data were modeled under a likelihood-based, "robust design" capture-recapture analytic framework. We explicitly modeled and estimated ecological parameters such as time-specific abundance, density, survival, recruitment, temporary emigration, and transience, using models that incorporated effects of factors such as individual heterogeneity, trap-response, and time on probabilities of photo-capturing tigers. The model estimated a random temporary emigration parameter of gamma" = gamma' = 0.10 +/- 0.069 (values are estimated mean +/- SE). When scaled to an annual basis, tiger survival rates were estimated at S = 0.77 +/- 0.051, and the estimated probability that a newly caught animal was a transient was tau = 0.18 +/- 0.11. During the period when the sampled area was of constant size, the estimated population size N(t) varied from 17 +/- 1.7 to 31 +/- 2.1 tigers, with a geometric mean rate of annual population change estimated as lambda = 1.03 +/- 0.020, representing a 3% annual increase. The estimated recruitment of new animals, B(t), varied from 0 +/- 3.0 to 14 +/- 2.9 tigers. Population density estimates, D, ranged from 7.33 +/- 0.8 tigers/100 km2 to 21.73 +/- 1.7 tigers/100 km2 during the study. Thus, despite substantial annual losses and temporal variation in recruitment, the tiger density remained at relatively high levels in Nagarahole. Our results are consistent with the hypothesis that protected wild tiger populations can remain healthy despite heavy mortalities because of their inherently high reproductive potential. The ability to model the entire photographic capture history data set and incorporate reduced-parameter models led to estimates of mean annual population change that were sufficiently precise to be useful. This efficient, noninvasive sampling approach can be used to rigorously investigate the population dynamics of tigers and other elusive, rare, wide-ranging animal species in which individuals can be identified from photographs or other means.

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.

Local hunting and the conservation of large mammals in India.

Hunting by local communities is among the most widespread threats to Indian wildlife, yet, the understanding of its nature, extent, and impacts on wildlife has been poor. We surveyed 2 protected areas--Kudremukha and Nagara-holé--in southern India to assess the impacts of local hunting on large mammals. Detailed interviews with retired and active hunters were employed to describe hunting patterns. Impacts of hunting were assessed by comparing large-mammal abundance in adjacent sites differing in their vulnerability to hunting. In Kudremukha, at least 26 species of mammals were hunted, mostly with guns, at an estimated intensity of 216 hunter-days per month per village. In Nagaraholé, 6 of the 9 focal species of large mammals occurred at significantly lower densities at the heavily hunted site where enforcement capabilities were poorer. Our data underscore the importance of preservationist programs in the conservation of large mammals in a context of extensive local hunting.

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.