Elevation and landscape change drive the distribution of a montane, endemic grassland bird.

CONTEXT: Tropical montane habitats support high biodiversity and are hotspots of endemism, with grasslands being integral components of many such landscapes. The montane grasslands of the Western Ghats have seen extensive land-use change over anthropogenic timescales. The factors influencing the ability of grassland-dependent species to persist in habitats experiencing loss and fragmentation, particularly in montane grasslands, are poorly known. OBJECTIVES: We studied the relationship between the Nilgiri pipit Anthus nilghiriensis, a threatened endemic bird that typifies these montane grasslands, and its habitat, across most of its global distribution. We examined what habitat features make remnant grasslands viable, which is necessary for their effective management. METHODS: We conducted 663 surveys in 170 sites and used both single-season occupancy modeling and N-mixture modeling to account for processes influencing detection, presence, and abundance. RESULTS: Elevation had a positive influence on species presence, patch size had a moderate positive influence, and patch isolation had a moderate negative influence. Species abundance was positively influenced by elevation and characteristics related to habitat structure, and negatively influenced by the presence of invasive woody vegetation. CONCLUSIONS: The strong effect of elevation on the highly range-restricted Nilgiri pipit is likely to make it vulnerable to climate change. This highly range-restricted species is locally extinct at several locations, and persists at low densities in remnants of its habitat left by recent fragmentation. Our findings indicate a need to control and reverse the spread of exotic woody invasives to preserve the grasslands themselves and the specialist species dependent upon them.

Author Correction: Towards a reliable assessment of Asian elephant population parameters: the application of photographic spatial capture-recapture sampling in a priority floodplain ecosystem.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

Towards a reliable assessment of Asian elephant population parameters: the application of photographic spatial capture-recapture sampling in a priority floodplain ecosystem.

The hitherto difficult task of reliably estimating populations of wide-ranging megafauna has been enabled by advances in capture-recapture methodology. Here we combine photographic sampling with a Bayesian spatially-explicit capture-recapture (SCR) model to estimate population parameters for the endangered Asian elephant Elephas maximus in the productive floodplain ecosystem of Kaziranga National Park, India. Posterior density estimates of herd-living adult females and sub-adult males and females (herd-adults) was 0.68 elephants/km2 (95% Credible Intervals, CrI = 0.56-0.81) while that of adult males was 0.24 elephants/km2 (95% CrI = 0.18-0.30), with posterior density estimates highlighting spatial heterogeneity in elephant distribution. Estimates of the space-usage parameter suggested that herd-adults ([Formula: see text] = 5.91 km, 95% CrI = 5.18-6.81) moved around considerably more than adult males ([Formula: see text] = 3.64 km, 95% CrI = 3.09-4.34). Based on elephant movement and age-sex composition, we derived the population that contributed individuals sampled in Kaziranga to be 908 herd-adults, 228 adult males and 610 young (density = 0.46 young/km2, SD = 0.06). Our study demonstrates how SCR is suited to estimating geographically open populations, characterising spatial heterogeneity in fine-scale density, and facilitating reliable monitoring to assess population status and dynamics for science and conservation.

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.

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.

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.