Niche Partitioning of 5 Sympatric Bornean Ungulates in Response to Spatiotemporal Variation in Fruit Availability and Resulting Implications for Predator-Prey Interactions

<p>Borneo’s rainforests are experiencing some of the fastest deforestation rates worldwide and are home to increasingly vulnerable species, most of which remain poorly understood. Bornean rainforests exhibit dramatic fluctuations in fruit and seed availability during mast-fruiting events which can exert considerable influence on frugivore ecology. Comprehensive spatiotemporal assessments of habitat use, resource partitioning, and responses to fruit availability in mast-fruiting rainforests are lacking for most species, including ungulates. The distribution and habitat use of an apex predator, the Sunda clouded leopard (Neofelis diardi), may be largely shaped by the availability of these ungulates. Yet, factors driving the spatial ecology of this elusive felid remain uncertain. I aimed to quantify spatiotemporal habitat use dynamics of these species and consequently inform effective conservation planning. Specifically, I quantified the effects of human activity, forest type, elevation, and mast-induced fluctuations in resources on the habitat use of lesser mousedeer (Tragulus kanchil), greater mousedeer (T. napu), Bornean yellow muntjacs (Muntiacus atherodes), red muntjacs (M. muntjak), and bearded pigs (Sus barbatus) in Gunung Palung National Park, West Kalimantan, Indonesian Borneo. I applied data from an extensive camera trapping study (n = 42,610 trap nights) to a modified single-season occupancy model to evaluate habitat use over space and time. I then applied estimates of occurrence (Ψ) of the five ungulate species to quantify if habitat use of the Sunda clouded leopard was influenced by prey occurrence and thus if this apex predator responded to bottom-up effects of resource variability. The results from the ungulate modelling revealed that forest type was an important predictor of habitat use of all ungulate species, each preferring different forest habitats. Habitat use estimates were highest in peat swamp forests for lesser mousedeer (Ψ = 0.92 ± 0.05), alluvial bench forests for greater mousedeer (Ψ = 0.52 ± 0.08), lowland granite forests for yellow (Ψ = 0.95 ± 0.07) and red muntjacs (Ψ = 0.98 ± 0.09), and freshwater swamp forests for bearded pigs (Ψ = 0.84 ± 0.07). Bearded pigs exhibited a link between variation in fruit availability and habitat use, indicating an ability to respond to resource variability. Occupancy modelling for Sunda clouded leopards revealed forest type, fruit availability, and bearded pig occurrence as the best predictors of habitat use. The highest estimates were associated with lowland granite forests (Ψ = 0.87 ± 0.09). My results reveal a novel pattern of niche partitioning through both food and habitat resources among five sympatric ungulate species and demonstrate that Sunda clouded leopards may use fruiting events as a cue for abundant prey. My research sheds light on important factors influencing habitat use of understudied ungulates and an apex predator and can be used to refine estimates of habitat suitability across a greater landscape to inform conservation practice amidst continually shrinking remnant forests in Indonesian Borneo.</p>

Niche Partitioning of 5 Sympatric Bornean Ungulates in Response to Spatiotemporal Variation in Fruit Availability and Resulting Implications for Predator-Prey Interactions

<p>Borneo’s rainforests are experiencing some of the fastest deforestation rates worldwide and are home to increasingly vulnerable species, most of which remain poorly understood. Bornean rainforests exhibit dramatic fluctuations in fruit and seed availability during mast-fruiting events which can exert considerable influence on frugivore ecology. Comprehensive spatiotemporal assessments of habitat use, resource partitioning, and responses to fruit availability in mast-fruiting rainforests are lacking for most species, including ungulates. The distribution and habitat use of an apex predator, the Sunda clouded leopard (Neofelis diardi), may be largely shaped by the availability of these ungulates. Yet, factors driving the spatial ecology of this elusive felid remain uncertain. I aimed to quantify spatiotemporal habitat use dynamics of these species and consequently inform effective conservation planning. Specifically, I quantified the effects of human activity, forest type, elevation, and mast-induced fluctuations in resources on the habitat use of lesser mousedeer (Tragulus kanchil), greater mousedeer (T. napu), Bornean yellow muntjacs (Muntiacus atherodes), red muntjacs (M. muntjak), and bearded pigs (Sus barbatus) in Gunung Palung National Park, West Kalimantan, Indonesian Borneo. I applied data from an extensive camera trapping study (n = 42,610 trap nights) to a modified single-season occupancy model to evaluate habitat use over space and time. I then applied estimates of occurrence (Ψ) of the five ungulate species to quantify if habitat use of the Sunda clouded leopard was influenced by prey occurrence and thus if this apex predator responded to bottom-up effects of resource variability. The results from the ungulate modelling revealed that forest type was an important predictor of habitat use of all ungulate species, each preferring different forest habitats. Habitat use estimates were highest in peat swamp forests for lesser mousedeer (Ψ = 0.92 ± 0.05), alluvial bench forests for greater mousedeer (Ψ = 0.52 ± 0.08), lowland granite forests for yellow (Ψ = 0.95 ± 0.07) and red muntjacs (Ψ = 0.98 ± 0.09), and freshwater swamp forests for bearded pigs (Ψ = 0.84 ± 0.07). Bearded pigs exhibited a link between variation in fruit availability and habitat use, indicating an ability to respond to resource variability. Occupancy modelling for Sunda clouded leopards revealed forest type, fruit availability, and bearded pig occurrence as the best predictors of habitat use. The highest estimates were associated with lowland granite forests (Ψ = 0.87 ± 0.09). My results reveal a novel pattern of niche partitioning through both food and habitat resources among five sympatric ungulate species and demonstrate that Sunda clouded leopards may use fruiting events as a cue for abundant prey. My research sheds light on important factors influencing habitat use of understudied ungulates and an apex predator and can be used to refine estimates of habitat suitability across a greater landscape to inform conservation practice amidst continually shrinking remnant forests in Indonesian Borneo.</p>

Density trends of wild felids in northern Laos

Determining the density trends of a guild of species can help illuminate their interactions, and the impacts that humans might have on them. We estimated the density trends from 2013 to 2017 of the clouded leopard Neofelis nebulosa, leopard cat Prionailurus bengalensis and marbled cat Pardofelis marmorata in Nam Et—Phou Louey National Park (NEPL), Laos, using camera trap data and spatial capture-recapture models. Mean (± SD) density estimates (individuals/100 km2) for all years were 1.77 ± 0.30 for clouded leopard, 1.50 ± 0.30 for leopard cat, and 3.80 ± 0.70 for marbled cat. There was a declining trend in density across the study years for all three species, with a ≥ 90% probability of decline for clouded leopard and leopard cat and an 83% probability of decline for marbled cat. There was no evidence that mesopredator release occurred as a result of tiger (Panthera tigris) and leopard (P. pardus) extirpations. We believe that snaring, the factor that led to the extirpation of tiger and leopard in NEPL, is now contributing to the decline of smaller felids, to an extent that over-rides any potential effects of mesopredator release on their densities and interactions. We recommend that the NEPL managers implement a more systematic and intensified snare removal program, in concert with extensive community outreach and engagement of local people to prevent the setting of snares. These actions might be the only hope for saving the remaining members of the felid community in NEPL.

The illegal exploitation of the Javan Leopard (Panthera pardus melas) and Sunda Clouded Leopard (Neofelis diardi) in Indonesia

Indonesia is home to the Javan Leopard (Panthera pardus melas) and the Sunda Clouded Leopard (Neofelis diardi), both of which are threatened by habitat loss, human-wildlife conflict issues and the illegal wildlife trade. Leopards and clouded leopards are threatened by the illegal wildlife trade across their range, however, very little is known of the illegal trade in these two species in Indonesia, or of the efforts made to tackle this crime. Both the Javan Leopard and Sunda Clouded Leopard are protected species in Indonesia and both species are listed in Appendix I of the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES), meaning commercial international trade is generally prohibited. To better understand the trade, and efforts to end this trade, we collected records of seizures and prosecutions relating to Javan Leopards and Sunda Clouded Leopards in Indonesia for the period 2011–2019. Despite both species being protected by law, this study reveals the prevalence of extensive poaching, illegal domestic trade and international trafficking of both species. A total of 41 seizure records were obtained from 2011 to 2019, which was estimated to amount to approximately 83 animals, which likely represents only a fraction of the total number of cases and therefore the risk may be substantially greater. Approximately half of the cases resulted in successful prosecution and of these, the highest sentence given was 2 years in jail and a fine of IDR50mil (~USD3300). The majority of the penalties handed down for these crimes were far below the maximum potential penalties and are unlikely to be effective deterrents.

Investigating carnivore guild structure: Spatial and temporal relationships amongst threatened felids in Myanmar

At least nine felid species can co-occur in Southeast Asia, thus providing an unusual opportunity to investigate poorly known guild structure and the factors controlling it. Using camera-trap data, we quantified space use, temporal activity, and multi-dimensional niche overlap of tiger, clouded leopard, marbled cat, leopard cat, and Asiatic golden cat in Htamanthi Wildlife Sanctuary of Myanmar. We hypothesized that the spatio-temporal behaviour of smaller cats can reflect avoidance of the larger cats, which are both potential competitors and predators, and similar-sized guild members would partition their niches in space or time to reduce competition for resources. Our approach involved single-species occupancy modelling to identify site covariates, pairwise spatial overlap using Bayesian inference, and activity overlap with Kernel density estimation and multivariate analyses to test hypotheses. We found tiger and marbled cats were primarily diurnal, clouded leopard and leopard cat were nocturnal and golden cat exhibited cathemeral activity. We observed a complex pattern of guild assembly and potential competition involving strong niche displacement between the golden cat and marbled cat, but high overlap between the relatively similarly-sized pairing of clouded leopard and golden cat, and the markedly differently-sized tiger – golden cat pairing. No significant evidence of mesopredator release was observed and the felid assembly in Northern Myanmar appeared to be partitioned mainly on a spatial, rather than temporal, dimension. Nonetheless, the temporal association between the three mesopredators was inversely related to the similarity in their body sizes. The insights into this felid guild revealed that the largest niche differences in the use of space and time occurred between the three smaller species, most evidently between the Asiatic golden cat-marbled cat pairing, followed by marbled cat - leopard cat pairing. This study offers new insight into carnivore guild assembly and, adds substantially to knowledge of five of the least known felids of conservation concern.

Clouded Leopard Neofelis nebulosa (Griffith, 1821) (Mammalia: Carnivora: Felidae) in illegal wildlife trade in Nepal

We document trade of the Clouded Leopard Neofelis nebulosa in Nepal based on pelt seizure reports published in wildlife trade reports and in newspapers. Just 27 cases in three decades seem little to suggest targeted illegal trade of the species, the seizure information in recent years indicate that illegal trade of Clouded Leopard body parts is still taking place. Hence an in-depth assessment is necessary to understand properly the intensity and magnitude of illegal trade on Clouded Leopard in the country.

Male residency of Sunda Clouded Leopard Neofelis diardi (Cuvier, 1823) (Mammalia: Carnivora: Felidae) in a peat swamp forest, Indonesian Borneo

The Sunda Clouded Leopard Neofelis diardi is the apex predator on the island of Borneo, yet little is known of its ecology. We document the length of residency of male Sunda Clouded Leopards in Central Kalimantan, Indonesian Borneo. Over 10 years, camera trap data have been obtained in the Sebangau peat swamp forest in a study area of ~ 105km2. We identified 11 individuals (eight males, one female, and two with unknown sex), from 152 notionally independent photographs. On average, males remained in the study area for 39.3 months (SE 8.3), or 3.3 years (SE 0.7), ranging from less than a month of residency up to 71 months. Females were infrequently recorded, possibly as a result of human disturbance and/or high male densities. Our results reveal that even a 10-year dataset is inadequate to answer some basic ecological questions, emphasising the importance of long-term monitoring of this species.

Temporal partitioning by felids, dholes and their potential prey in northern Laos

Temporal partitioning can allow sympatric carnivores to coexist, especially if overlap of other resources is high. Using camera trap data from 2013 to 2017, we investigated the temporal partitioning of a community of wild felids and a canid in Nam Et–Phou Louey National Protected Area, Laos, to determine the extent to which temporal avoidance might be facilitating coexistence of similarly sized carnivores. We also investigated temporal overlap of these carnivore species and their presumed main prey, to determine if their activity is likely most influenced by their prey or potential competitors. The dhole (Cuon alpinus) and clouded leopard (Neofelis nebulosa), the two largest carnivores, had low temporal overlap, and activity appeared to be synchronized with the main prey for dhole, but not clouded leopard. Thus, it was possible that clouded leopard used temporal partitioning to coexist with dhole. The temporal overlap of Asian golden cat (Catopuma temminckii) was high with clouded leopard and moderate with dhole, and overall appeared to be influenced most by its presumed prey species, rather than by its potential competitors. The two small felids had the least temporal overlap between any species, as the marbled cat (Pardofelis marmorata) was primarily diurnal whereas the leopard cat (Prionailurus bengalensis) was primarily nocturnal. Given that the two small felids reportedly have similar diets consisting of small rodents and birds, their temporal partitioning is likely to help facilitate their coexistence. For carnivore species in northern Laos that are most similar in body size and diet, temporal partitioning is likely to be an important mechanism of coexistence. Otherwise, temporal patterns appear to be synchronized with their main prey.