Andreas Wilting

Modelling the species distribution of flat-headed cats (Prionailurus planiceps), an endangered South-East Asian small felid.

Background The flat-headed cat (Prionailurus planiceps) is one of the worlds least known, highly threatened felids with a distribution restricted to tropical lowland rainforests in Peninsular Thailand/Malaysia, Borneo and Sumatra. Throughout its geographic range large-scale anthropogenic transformation processes, including the pollution of fresh-water river systems and landscape fragmentation, raise concerns regarding its conservation status. Despite an increasing number of camera-trapping field surveys for carnivores in South-East Asia during the past two decades, few of these studies recorded the flat-headed cat. Methodology/Principal Findings In this study, we designed a predictive species distribution model using the Maximum Entropy (MaxEnt) algorithm to reassess the potential current distribution and conservation status of the flat-headed cat. Eighty-eight independent species occurrence records were gathered from field surveys, literature records, and museum collections. These current and historical records were analysed in relation to bioclimatic variables (WorldClim), altitude (SRTM) and minimum distance to larger water resources (Digital Chart of the World). Distance to water was identified as the key predictor for the occurrence of flat-headed cats (>50% explanation). In addition, we used different land cover maps (GLC2000, GlobCover and SarVision LLC for Borneo), information on protected areas and regional human population density data to extract suitable habitats from the potential distribution predicted by the MaxEnt model. Between 54% and 68% of suitable habitat has already been converted to unsuitable land cover types (e.g. croplands, plantations), and only between 10% and 20% of suitable land cover is categorised as fully protected according to the IUCN criteria. The remaining habitats are highly fragmented and only a few larger forest patches remain. Conclusion/Significance Based on our findings, we recommend that future conservation efforts for the flat-headed cat should focus on the identified remaining key localities and be implemented through a continuous dialogue between local stakeholders, conservationists and scientists to ensure its long-term survival. The flat-headed cat can serve as a flagship species for the protection of several other endangered species associated with the threatened tropical lowland forests and surface fresh-water sources in this region.

Density of the Vulnerable Sunda clouded leopard Neofelis diardi in two commercial forest reserves in Sabah, Malaysian Borneo

Recently the Sunda clouded leopard Neofelis diardi was recognized as a separate species distinct from the clouded leopard Neofelis nebulosa of mainland Asia. Both species are categorized as Vulnerable on the IUCN Red List. Little is known about the newly identified species and, in particular, information from forests outside protected areas is scarce. Here we present one of the first density estimates calculated with spatial capture–recapture models using camera-trap data. In two commercial forest reserves in Sabah (both certified for their sustainable management practices) the density of the Sunda clouded leopard was estimated to be c. 1 per 100 km 2 (0.84±SE 0.42 and 1.04±SE 0.58). The presence of the Sunda clouded leopard in such forests is encouraging for its conservation but additional studies from other areas, including protected forests, are needed to compare and evaluate these densities.

Targeted Conservation to Safeguard a Biodiversity Hotspot from Climate and Land-Cover Change

Responses of biodiversity to changes in both land cover and climate are recognized [1] but still poorly understood [2]. This poses significant challenges for spatial planning as species could shift, contract, expand, or maintain their range inside or outside protected areas [2-4]. We examine this problem in Borneo, a global biodiversity hotspot [5], using spatial prioritization analyses that maximize species conservation under multiple environmental-change forecasts. Climate projections indicate that 11%-36% of Bornean mammal species will lose ≥ 30% of their habitat by 2080, and suitable ecological conditions will shift upslope for 23%-46%. Deforestation exacerbates this process, increasing the proportion of species facing comparable habitat loss to 30%-49%, a 2-fold increase on historical trends. Accommodating these distributional changes will require conserving land outside existing protected areas, but this may be less than anticipated from models incorporating deforestation alone because some species will colonize high-elevation reserves. Our results demonstrate the increasing importance of upland reserves and that relatively small additions (16,000-28,000 km(2)) to the current conservation estate could provide substantial benefits to biodiversity facing changes to land cover and climate. On Borneo, much of this land is under forestry jurisdiction, warranting targeted conservation partnerships to safeguard biodiversity in an era of global change.

Geographical variation in and evolutionary history of the Sunda clouded leopard (Neofelis diardi) (Mammalia: Carnivora: Felidae) with the description of a new subspecies from Borneo

Recent morphological and molecular studies led to the recognition of two extant species of clouded leopards; Neofelis nebulosa from mainland southeast Asia and Neofelis diardi from the Sunda Islands of Borneo and Sumatra, including the Batu Islands. In addition to these new species-level distinctions, preliminary molecular data suggested a genetic substructure that separates Bornean and Sumatran clouded leopards, indicating the possibility of two subspecies of N. diardi. This suggestion was based on an analysis of only three Sumatran and seven Bornean individuals. Accordingly, in this study we re-evaluated this proposed subspecies differentiation using additional molecular (mainly historical) samples of eight Bornean and 13 Sumatran clouded leopards; a craniometric analysis of 28 specimens; and examination of pelage morphology of 20 museum specimens and of photographs of 12 wild camera-trapped animals. Molecular (mtDNA and microsatellite loci), craniomandibular and dental analyses strongly support the differentiation of Bornean and Sumatran clouded leopards, but pelage characteristics fail to separate them completely, most probably owing to small sample sizes, but it may also reflect habitat similarities between the two islands and their recent divergence. However, some provisional discriminating pelage characters are presented that need further testing. According to our estimates both populations diverged from each other during the Middle to Late Pleistocene (between 400 and 120 kyr). We present a discussion on the evolutionary history of Neofelis diardi sspp. on the Sunda Shelf, a revised taxonomy for the Sunda clouded leopard, N. diardi, and formally describe the Bornean subspecies, Neofelis diardi borneensis, including the designation of a holotype (BM.3.4.9.2 from Baram, Sarawak) in accordance with the rules of the International Code of Zoological Nomenclature.

Density and habitat use of the leopard cat (Prionailurus bengalensis) in three commercial forest reserves in Sabah, Malaysian Borneo

Abstract The small (2- to 7-kg) leopard cat (Prionailurus bengalensis) is the most common cat species in Asia. Although it occurs in a wide range of habitats and seems to adapt well to anthropogenic habitat changes, surprisingly little is known about this species in the wild. All studies have focused on protected areas, although a large proportion of Southeast Asian forests are timber concessions. During this study, we used large camera-trapping data sets (783 records of 124 individuals) from 3 commercially used forests to investigate consequences of different logging regimes on density and habitat associations of the leopard cat. We applied spatial capture–recapture models accounting for the location of camera-traps (on or off road) to obtain estimates of leopard cat density. Density was higher in the 2 more disturbed forest reserves (X̄ = 12.4 individuals/100 km2 ± 1.6 SE and 16.5 ± 2 individuals/100 km2) than in the sustainably managed forest (9.6 ± 1.7 individuals/100 km2). Encounter rates with off-road traps were only 3.6–9.1% of those for on-road traps. Occupancy models, which accounted for spatial autocorrelation between sampling sites by using a conditional autoregressive model, revealed that canopy closure and ratio of climax to pioneer trees had a significantly negative impact on leopard cat occurrence. Our results confirm that the leopard cat is doing well in modified landscapes and even seems to benefit from the opening of forests. With such flexibility the leopard cat is an exception among tropical rain-forest carnivores.

Anticipated climate and land-cover changes reveal refuge areas for Borneo's orang-utans.

Habitat loss and climate change pose a double jeopardy for many threatened taxa, making the identification of optimal habitat for the future a conservation priority. Using a case study of the endangered Bornean orang-utan, we identify environmental refuges by integrating bioclimatic models with projected deforestation and oil-palm agriculture suitability from the 1950s to 2080s. We coupled a maximum entropy algorithm with information on habitat needs to predict suitable habitat for the present day and 1950s. We then projected to the 2020s, 2050s and 2080s in models incorporating only land-cover change, climate change or both processes combined. For future climate, we incorporated projections from four model and emission scenario combinations. For future land cover, we developed spatial deforestation predictions from 10 years of satellite data. Refuges were delineated as suitable forested habitats identified by all models that were also unsuitable for oil palm - a major threat to tropical biodiversity. Our analyses indicate that in 2010 up to 260,000 km(2) of Borneo was suitable habitat within the core orang-utan range; an 18-24% reduction since the 1950s. Land-cover models predicted further decline of 15-30% by the 2080s. Although habitat extent under future climate conditions varied among projections, there was majority consensus, particularly in north-eastern and western regions. Across projections habitat loss due to climate change alone averaged 63% by 2080, but 74% when also considering land-cover change. Refuge areas amounted to 2000-42,000 km(2) depending on thresholds used, with 900-17,000 km(2) outside the current species range. We demonstrate that efforts to halt deforestation could mediate some orang-utan habitat loss, but further decline of the most suitable areas is to be expected given projected changes to climate. Protected refuge areas could therefore become increasingly important for ongoing translocation efforts. We present an approach to help identify such areas for highly threatened species given environmental changes expected this century.

iDNA from terrestrial haematophagous leeches as a wildlife surveying and monitoring tool – prospects, pitfalls and avenues to be developed

Invertebrate-derived DNA (iDNA) from terrestrial haematophagous leeches has recently been proposed as a powerful non-invasive tool with which to detect vertebrate species and thus to survey their populations. However, to date little attention has been given to whether and how this, or indeed any other iDNA-derived data, can be combined with state-of-the-art analytical tools to estimate wildlife abundances, population dynamics and distributions. In this review, we discuss the challenges that face the application of existing analytical methods such as site-occupancy and spatial capture-recapture (SCR) models to terrestrial leech iDNA, in particular, possible violations of key assumptions arising from factors intrinsic to invertebrate parasite biology. Specifically, we review the advantages and disadvantages of terrestrial leeches as a source of iDNA and summarize the utility of leeches for presence, occupancy, and spatial capture-recapture models. The main source of uncertainty that attends species detections derived from leech gut contents is attributable to uncertainty about the spatio-temporal sampling frame, since leeches retain host-blood for months and can move after feeding. Subsequently, we briefly address how the analytical challenges associated with leeches may apply to other sources of iDNA. Our review highlights that despite the considerable potential of leech (and indeed any) iDNA as a new survey tool, further pilot studies are needed to assess how analytical methods can overcome or not the potential biases and assumption violations of the new field of iDNA. Specifically we argue that studies to compare iDNA sampling with standard survey methods such as camera trapping, and those to improve our knowledge on leech (and other invertebrate parasite) physiology, taxonomy, and ecology will be of immense future value.

camtrapR: an R package for efficient camera trap data management

Summary Camera trapping is a widely applied method to study mammalian biodiversity and is still gaining popularity. It can quickly generate large amounts of data which need to be managed in an efficient and transparent way that links data acquisition with analytical tools. We describe the free and open-source R package camtrapR, a new toolbox for flexible and efficient management of data generated in camera trap-based wildlife studies. The package implements a complete workflow for processing camera trapping data. It assists in image organization, species and individual identification, data extraction from images, tabulation and visualization of results and export of data for subsequent analyses. There is no limitation to the number of images stored in this data management system; the system is portable and compatible across operating systems. The functions provide extensive automation to minimize data entry mistakes and, apart from species and individual identification, require minimal manual user input. Species and individual identification are performed outside the R environment, either via tags assigned in dedicated image management software or by moving images into species directories. Input for occupancy and (spatial) capture–recapture analyses for density and abundance estimation, for example in the R packages unmarked or secr, is computed in a flexible and reproducible manner. In addition, survey summary reports can be generated, spatial distributions of records can be plotted and exported to gis software, and single- and two-species activity patterns can be visualized. camtrapR allows for streamlined and flexible camera trap data management and should be most useful to researchers and practitioners who regularly handle large amounts of camera trapping data.

Coming down from the trees: Is terrestrial activity in Bornean orangutans natural or disturbance driven?

The orangutan is the worlds largest arboreal mammal, and images of the red ape moving through the tropical forest canopy symbolise its typical arboreal behaviour. Records of terrestrial behaviour are scarce and often associated with habitat disturbance. We conducted a large-scale species-level analysis of ground-based camera-trapping data to evaluate the extent to which Bornean orangutans Pongo pygmaeus come down from the trees to travel terrestrially, and whether they are indeed forced to the ground primarily by anthropogenic forest disturbances. Although the degree of forest disturbance and canopy gap size influenced terrestriality, orangutans were recorded on the ground as frequently in heavily degraded habitats as in primary forests. Furthermore, all age-sex classes were recorded on the ground (flanged males more often). This suggests that terrestrial locomotion is part of the Bornean orangutans natural behavioural repertoire to a much greater extent than previously thought, and is only modified by habitat disturbance. The capacity of orangutans to come down from the trees may increase their ability to cope with at least smaller-scale forest fragmentation, and to cross moderately open spaces in mosaic landscapes, although the extent of this versatility remains to be investigated.

Connecting Earth observation to high-throughput biodiversity data

Understandably, given the fast pace of biodiversity loss, there is much interest in using Earth observation technology to track biodiversity, ecosystem functions and ecosystem services. However, because most biodiversity is invisible to Earth observation, indicators based on Earth observation could be misleading and reduce the effectiveness of nature conservation and even unintentionally decrease conservation effort. We describe an approach that combines automated recording devices, high-throughput DNA sequencing and modern ecological modelling to extract much more of the information available in Earth observation data. This approach is achievable now, offering efficient and near-real-time monitoring of management impacts on biodiversity and its functions and services.