|Scientific Name:||Neofelis diardi|
|Species Authority:||(G. Cuvier, 1823)|
|Infra-specific Taxa Assessed:|
|Taxonomic Notes:||Classically considered a single species, the Clouded Leopard has been split into two species. Based on analysis of mitochondrial DNA, microsatellites, chromosomal differences and skull and pelage characteristics, Neofelis nebulosa is restricted to mainland southeast Asia, and N. diardi, the Sunda Clouded Leopard, is found on the islands of Sumatra and Borneo (Buckley-Beason et al. 2006; Kitchener et al. 2006, 2007; Christiansen et al. 2008, 2009; Wilting et al. 2007; Wilting et al. 2011). Sundaland or the Sunda regions refer to the islands of Sumatra, Borneo, Java and Bali as well as to the Malay Peninsula. Although samples are limited from Peninsular Malaysia, it appears to be inhabited by the mainland Clouded Leopard N. nebulosa, rather than the Sunda Island type (Kitchener et al. 2007). The original type specimen of N. diardi originated from Sumatra, but it was initially recorded in error as from Java (Kitchener et al. 2006). As this type specimen described by Cuvier (1823) is not present in any museum, Christiansen 2009 designated a Neoptype (RMNH1981) from Palembang, Sumatra, which is housed at the National Museum of Natural History (Naturalis), in Leiden, the Netherlands. Clouded Leopard fossils have been found on Java, where it perhaps became extinct in the Holocene (Meijaard 2004). Based on analysis of genetic, skull and pelage characteristics, Wilting et al. (2011) recognized two distinct subspecies of N. diardi: the Sumatran Clouded Leopard N. d. diardi and the Bornean Clouded Leopard N. d. borneensis.|
|Red List Category & Criteria:||Vulnerable A2c; C1 ver 3.1|
|Assessor(s):||Hearn, A., Ross, J., Brodie, J., Cheyne, S., Haidir, I.A., Loken, B., Mathai, J., Wilting, A. & McCarthy, J.|
|Reviewer(s):||Nowell, K., Hunter, L., Duckworth, J.W., Breitenmoser-Würsten, C., Lanz, T. & Breitenmoser, U.|
|Contributor(s):||Sanderson, J. & Sunarto, S.|
The elusive Sunda Clouded Leopard is found only on the islands of Borneo and Sumatra, and possibly on the Batu Islands. Although thought to exist at relatively low population densities, the Sunda Clouded Leopard appears to be adaptable, and is found in a range of forest types, elevations and levels of disturbance. Despite the species’ apparent habitat plasticity, the Sunda Clouded Leopard is forest dependent, and does not readily use oil palm Elaeis guineensis plantations (Ross et al. 2010, Hearn, Ross and Macdonald unpublished data), although such plantations may not necessarily form a hard barrier to this wild cat (Nájera et al. 2013, Hearn et al. unpublished data). The forests of Borneo and Sumatra are undergoing some of the world’s highest deforestation rates, largely as the result of the expansion of oil palm plantations, and thus such development and consequent loss of habitat, coupled with the species’ apparent low population size, probably constitute the greatest threat to this species.
Population density estimates of Sunda Clouded Leopard on Borneo, derived from targeted camera trap surveys across a range of forest types and elevations, and a gradient of disturbance, range from 0.8 to 4.4 individuals / 100 km² (Brodie and Giordano 2012, Wilting et al. 2012, Cheyne and Macdonald 2011, Hearn, Ross and Macdonald unpublished data, Loken et al. in prep), a large proportion of which (6 of 10 surveys) provide estimates of under 1.9 individuals / 100 km². Additional camera trap surveys on Borneo, specifically undertaken to detect and estimate the density of this felid, failed to find any evidence of the Sunda Clouded Leopard, including the Kabili-Sepilok Forest Reserve, Sabah (Hearn, Ross and Macdonald unpublished data) and several forest sites in Kalimantan (Cheyne et al. in prep), indicating that Sunda Clouded Leopard presence across the forested landscape of Borneo is somewhat patchy. The population density of the Sunda Clouded Leopard on Sumatra is less well studied, but four camera surveys undertaken within the Kerinci landscape present a similar pattern to that of Borneo, with densities ranging from 0.8 to 1.6 individuals / 100 km² (Sollmann et al. 2014).
A Maximum Entropy based habitat suitability analysis, using presence-only and expert opinion data in combination with environmental and land use covariates, estimated that the Sunda Clouded Leopard’s current area of occupancy (AOO) on Borneo is approximately 378,900 km² (Borneo Carnivore Symposium: Hearn et al. in prep). Such habitat suitability data are lacking for Sumatra. For this assessment, an approximate AOO for Sumatra was developed from a map of forest cover derived from satellite images taken in 2010 (Miettinen et al. 2010). Land cover classes identified as suitable habitat by expert opinion during the Bornean Carnivore Symposium in 2011 (Peat-swamp forest, Lowland Forest and lower montane forest <1500 m) were selected and included in the AOO, which amounted to approximately 73,000 km², providing a total AOO of 451,900 km². As detailed below, forest cover on both Borneo and Sumatra will have decreased since 2010. Furthermore, due to a lack of sufficient data, these estimates are unable to account for the potential impacts of differential hunting pressure on population density throughout this range, which could plausibly be having a considerable impact on the species and our distribution estimate may, therefore, represent an optimistic estimation of the current AOO. Acknowledging that our estimate of current AOO may be biased high as a result of unmeasured hunting pressure and a patchy distribution, and further loss of forest post 2010, we take a precautionary approach and estimate that the mean density of the Sunda Clouded Leopard across its AOO may be as low as 1 individual / 100 km². Extrapolation of this estimate to the wider AOO suggests that the number of mature individuals could be around 4,500 individuals, and is thus very likely to be below 10,000 individuals.
A GIS exercise as part of the current assessment used the habitat suitability assessment alongside land cover data for Borneo for 2000 and 2010 (Miettinen et al. 2010), and the selected land cover classes for Sumatra for the years 2000 and 2010 (Miettinen et al. 2010) and estimated that the AOO decreased by 33.1% during this time period, primarily as a result of forest loss and conversion to oil palm plantations. Given the strong evidence for forest dependency in this felid, this loss of AOO, coupled with hunting pressure, is probably driving a reduction in the number of mature individuals at the same rate or higher. More recent satellite imagery shows that forest loss has continued to reduce the AOO (Gaveau et al. 2014, Macdonald et al. in prep a), and it is likely that forest loss continues to this day, albeit at a reduced rate. Although the Sunda Clouded Leopard probably spends a proportion of its time hunting, moving and resting in the trees, because much of its time is also spent moving on the ground, Sunda Clouded Leopards are prone to untargeted snaring.
We therefore have a strong basis upon which to infer that the total Sunda Clouded Leopard population size currently lies below 10,000 individuals, and that the cumulative reduction in the AOO and thus population size as a consequence of forest loss, degradation and poaching has probably amounted to ≥30% in the past 21 years (three Sunda Clouded Leopard generations, Pacifici et al. 2013) indicating a categorisation of Vulnerable under both VU C1 and VU A2c.
|Previously published Red List assessments:||
|Range Description:||The Sunda Clouded Leopard is found on the islands of Sumatra and Borneo (Buckley-Beason et al. 2006; Kitchener et al. 2006, 2007; Christiansen et al. 2008; Christiansen 2009; Wilting et al. 2007, 2011). It is unknown if there are still Sunda Clouded Leopards on the small Batu Islands close to Sumatra. There are Clouded Leopard fossils from Java (Meijaard 2004), but no records of living animals from modern times. A Maximum Entropy habitat suitability model for Sunda Clouded Leopards on Borneo, derived from presence-only and expert opinion data in combination with environmental and land use covariates (Borneo Carnivore Symposium: Hearn et al. in prep), was used to develop an estimate of this species’ current area of occupancy (AOO) on Borneo, and provided an estimate of approximately 378,900 km². Data regarding Sunda Clouded Leopard habitat suitability are currently unavailable for Sumatra, however, and so as part of this assessment, an approximate AOO for Sumatra was developed by quantifying areas of suitable habitat from a land cover map derived from satellite images taken in 2010 (Miettinen et al. 2012), based on land cover classes identified as suitable habitat by expert opinion during the Bornean Carnivore Symposium in 2011. The analysis provided an approximate AOO for Sumatra of 73,000 km², and thus a range-wide total AOO of 451,900 km². As detailed below, forest cover on both Borneo and Sumatra will have decreased since 2010. Furthermore, due to a lack of sufficient data, these estimates are unable to account for the potential impacts of differential hunting pressure on population density throughout this range, which could plausibly be having a considerable impact on the species and our distribution estimate may, therefore, represent an optimistic estimation of the current AOO.
The habitat suitability analysis for Borneo suggest that this felid has a widespread distribution over a large contiguous forested portion of Borneo, with the only exception being South Kalimantan, which was predicted as being largely composed of non-habitat. The distribution map presented as part of this assessment is largely based on this habitat suitability exercise, but small patches which were identified as non-habitat in the model output have been included here as habitat. In addition, the Tabin Wildlife Reserve, not included in the original model output, has been included here, based on recorded presence (Ross et al. 2010, Bernard et al. 2012).
Native:Brunei Darussalam; Indonesia (Kalimantan, Sumatera); Malaysia (Sabah, Sarawak)
|Upper elevation limit (metres):||1500|
|Range Map:||Click here to open the map viewer and explore range.|
|Population:||Our understanding of the Sunda Clouded Leopard’s population density on Borneo, and particularly in the Malaysian state of Sabah, has improved in recent years, with the advent of intensive, targeted camera traps surveys, analysed within a capture-recapture framework. Population density estimates from a range of forest types and elevations, and a gradient of disturbance are typically low (0.8 – 4.4 ind. / 100 km², with 6 of 10 surveys providing estimates of under 1.9 ind. / 100 km². All estimates from Sabah were derived using a spatially explicit capture recapture framework (SECR), from primary Dipterocarp forest, including lowland (Maliau Basin Conservation Area, 1.9 ind. / 100 km²: Brodie and Giordano 2012, Danum Valley Conservation Area, 1.76 ind. / 100 km²: Hearn, Ross and Macdonald unpublished data), upland (Tawau Hills National Park, 1.85 ind. / 100 km²: Hearn, Ross and Macdonald unpublished data) and submontane/montane (Crocker Range National Park, 1.4 ind. / 100 km²: Hearn, Ross and Macdonald unpublished data), population density estimates from selectively logged forests are more variable (Tangkulap-Pinangah Forest Reserve, 0.84 ind. / 100 km2, Segaliud Lokan Forest Reserve, 1.0 ind. / 100 km²: Wilting et al. 2012, Ulu Segama Forest Reserve, 2.55 ind. / 100 km², Tabin Wildlife Reserve, 2.26 ind. / 100 km² (Hearn, Ross and Macdonald unpublished data). In Kalimantan, there are only two studies that have yielded population density estimates of the Sunda Clouded Leopard. Loken et al. (in prep) used SECR and provide estimates from the selectively logged Wehea Forest in East Kalimantan (1.23 to 3.01 ind. / 100 km²). Using a conventional capture recapture framework, which typically results in higher densities from that derived using SECR, Cheyne et al. (2013) provide density estimates for a population of Sunda Clouded Leopards residing within the Sabangau peat swamp, a disturbed peat swamp forest in Central Kalimantan (0.7-4.4 ind. / 100 km²). Additional Sunda Clouded Leopard focused surveys on Borneo failed to find any evidence of this species, including the relatively small (ca. 43 km²) and isolated Kabili-Sepilok Forest Reserve, Sabah (Ross et al. 2013, Hearn, Ross and Macdonald unpublished data) and several forest sites in Kalimantan (Cheyne et al. in prep). Such surveys highlight the potential patchy nature of Sunda Clouded Leopard presence across the forested landscape of Borneo. The population density of this felid on Sumatra has received less attention, but surveys are currently underway (I. Haidir pers comm, 2015), and four camera surveys undertaken within the Kerinci landscape present a similar pattern to that of Borneo, with densities ranging from 0.8 to 1.6 individuals / 100 km² (Sollmann et al. 2014).
A Maximum Entropy based habitat suitability analysis for Borneo, discussed in detail above, provided a total AOO of 451,900 km² (Borneo: 378,900 km², Sumatra: 73,000 km²). Acknowledging that our estimate of current AOO may be biased high as a result of unmeasured hunting pressure, a potentially patchy distribution, and further loss of forest post-2010, we take a precautionary approach and estimate that the mean density of the Sunda Clouded Leopard across its AOO may plausibly be as low as 1 individual / 100 km². Extrapolation of this estimate to the wider AOO suggests that the number of mature individuals range-wide could be around 4,500 individuals, with around 3,800 persisting in Borneo and as few as 730 individuals remaining in Sumatra.
|Current Population Trend:||Decreasing|
|Habitat and Ecology:||The Sunda Clouded Leopard appears to be a relatively adaptable species, and is found in a range of forest types, elevations and levels of disturbance. Recent camera trap surveys have recorded the felid in primary lowland, upland and submontane Dipterocarp forest (Ross et al. 2010, Brodie and Giordano 2012, McCarthy et al. 2015, Loken et al. unpublished data, Hearn, Ross and Macdonald unpublished data), selectively logged Dipterocarp forest (Ross et al. 2010, Wilting et al. 2012, Mathai et al. 2014, Sollmann et al. 2014, Loken et al. unpublished data), and peat-swamp forest (Cheyne et al. 2011, 2013). Although the species is able to persist in modified habitats, the effects of anthropogenic disturbance remain unclear. Brodie et al. (2015) estimated the local abundance of Sunda Clouded Leopard through hierarchical modelling of camera-trap data with biophysical and anthropogenic covariates and found that an increased road density was associated with reduced local abundance of Sunda Clouded Leopards. Their study also showed that local abundance increased linearly as elevation increased, although they highlight that the effect was slight. Similar responses were shown by McCarthy et al. (2015), who applied a Maximum entropy approach to model habitat use of the Sunda Clouded Leopard in Bukit Barisan Selatan National Park, Sumatra, and reported the species appeared to be relatively intolerant of anthropogenic features, occurring most commonly at moderate distances from roads, rivers and forest edges, all features which assist the movement of people. McCarthy et al. (2015) also found a positive association with moderate elevation forest. Although there is much to learn about the response of the Sunda Clouded Leopard to habitat modification, the available evidence strongly supports the notion that this felid is forest dependent, and does not readily use oil palm Elaeis guineensis plantations (Ross et al. 2010, Yue et al. in press, Hearn, Ross and Macdonald unpublished data), although such plantations may not necessarily form a hard barrier to this wild cat (Nájera et al. 2013, Hearn et al. unpublished data).
On Sumatra, the Sunda Clouded Leopard was hypothesized to exist at much lower population densities than on Borneo, potentially in response to predation and/or interspecific competition from tigers. The few density estimates that have been recorded from Sumatra (Sollmann et al. 2014) to date, however, lie within the range of estimates for Borneo (Brodie and Giordano 2012, Wilting et al. 2012, Cheyne et al. 2013, Hearn, Ross and Macdonald unpublished data). Further data are thus needed to investigate whether interspecific competition is influencing densities of the Sunda Clouded Leopard.
Little is known regarding the spatial ecology of the Sunda Clouded Leopard. A female Sunda Clouded Leopard, residing in a selectively logged forest in Sabah, occupied a home-range of 16.1 km² and a core-range of 5.4 km² (95% and 50% fixed kernel estimators, respectively) (Hearn et al. 2013). The data only spanned a 109 day period, however, yet they represent the only available published data on the movements of this species. Clouded Leopards (Neofelis spp.) possess morphological adaptions that enable them to climb and move swiftly in the forest canopy, and there are numerous incidental records of such behaviour in the Sunda Clouded Leopard on Borneo (eg. Davis 1962, Matsuda et al. 2008, Morino 2010). Track surveys (Holden 2001, Gordon and Stewart 2007) and camera trap records (Ridout and Linkie 2009, Ross et al. 2010, Cheyne and Macdonald 2011, Brodie and Giordano 2012, Wilting et al. 2012, Cheyne et al. 2013, Hearn et al. 2013, Pusparini et al. 2014, McCarthy et al. 2015) highlight that terrestrial activity is commonplace, particularly along trails and logging roads. Rabinowitz et al. (1987) proposed that Clouded Leopards may be less arboreal on Borneo due to the absence of sympatric tigers. Movement data from tagged animals in Sabah show speeds which are indicative of a predominantly terrestrial activity pattern (Hearn et al. unpublished data).
No studies as yet have quantitatively investigated the diet of the Sunda Clouded Leopard. Observations of predation events, however, suggest that the species feeds on a wide range of small to medium sized mammalian prey, including Sambar Deer Rusa unicolor, Muntjacs Muntiacus spp., Bearded Pig Sus barbatus, Mouse Deer Tragulus spp., porcupines and primates, and it is likely that the diet of this felid varies considerably across its range (Davis 1962, Rabinowitz et al. 1987, Yeager 1991, Gordon and Stewart 2007, Matsuda et al. 2008, Mohamed et al. 2009, Morino 2010). Ross et al. (2013) used camera trap data to show that Bearded Pigs exhibited greater nocturnal activity in the absence of Sunda Clouded Leopardc, with female pigs and young showing the strongest response. The study suggests that Bearded Pigs, and particularly young animals, may be a key prey species and are capable of altering their activity pattern in response to this predation risk.
Analyses of camera trap photographs from both Borneo (Ross et al. 2010, Cheyne and Macdonald 2011, Hearn et al. 2013) and Sumatra (Ridout and Linkie 2009, Pusparini et al. 2014, McCarthy et al. 2015) suggest the Sunda Clouded Leopard’s temporal activity is largely nocturnal, with significant activity being shown during crepuscular periods, and this pattern is supported by activity data from radio-collared animals in Sabah (Hearn et al. unpublished data). Such sources provide several instances of diurnal activity, however, and there are incidental records of Sunda Clouded Leopards hunting primates during the day (Davis 1962, Matsuda et al. 2008, Morino 2010).
|Generation Length (years):||7|
|Use and Trade:||See Threats section.|
Habitat loss due to commercial logging and conversion to oil palm plantations and the increasing threat from poaching pose the greatest threats to the Sunda Clouded Leopard. Oil palm plantations are likely to expand in the future as a result of the push for biofuels, and forest cover on the island of Borneo, if current deforestation rates continue, is projected to decline from 50% to less than one-third by 2020 (Rautner 2005). A more recent analysis of deforestation risk predicts that by 2020 37% of the remaining forest in Borneo will have a greater than 25% risk of being deforested and that approximately 13% of Borneo’s forests (an area of 11,167 km²) will have a greater than 50% chance of being lost (Macdonald et al. in prep. a). This deforestation will not only reduce the area of available habitat for the Sunda Clouded Leopard but it is also likely to increase the fragmentation of its populations. In a modelling analysis of the likely impacts of this deforestation, Macdonald et al. (in prep. b) predict that between 2010 and 2020 the percentage of the landscape connected by dispersal in Borneo will decrease by over 60% and that this could lead to both a large reduction in the Sunda Clouded Leopard numbers and also a significant reduction in the potential gene flow between populations. Increased fragmentation of Clouded Leopard habitat may also be resulting in an increased risk of disease in some populations due to exposure to pathogens present in feral cat and dog populations. One study in Sabah found seropositive findings for canine parvovirus and/or distemper virus in feral cats and dogs, Malay Civets (Viverra tangalunga) and Common Palm Civets (Paradoxurus hermaphroditus) living sympatrically with Sunda Clouded Leopards (Nájera et al. in prep).
Poaching, either directly through the use of snares, or through the reduction of prey availability by the poaching of game species, poses a significant threat, both in Sumatra (Holden et al. 2001) and Borneo (Rabinowitz et al. 1987, Wilting et al. 2006, Cheyne et al. 2013).
|Conservation Actions:||The Sunda Clouded Leopard is currently listed under CITES Appendix I (as Neofelis nebulosa), and warrants full listing under N. diardi. The species is fully protected in Sumatra and Kalimantan (Indonesia), Sabah and Sarawak (Malaysia), and Brunei. It occurs in most protected areas along the Sumatran mountain spine, and in most of the larger protected areas on Borneo, though maintaining or re-establishing connectivity among protected areas (e.g. via habitat corridors) remains a critical conservation priority for the species. The Sunda Clouded Leopard has been the subject of increased research in recent years, but it remains one of the least studied of the world’s Pantherine felids, hampering the development of conservation actions. It is essential, therefore, to gain an understanding of this species’ distribution, abundance, and responses to anthropogenic modification, particularly in Sumatra, and also in Kalimantan, Sarawak and Brunei. Data on the distribution and abundance of this species in Sabah has improved greatly in recent years, and it remains essential to monitor the population over time. A detailed study of the Sunda Clouded Leopard’s basic ecology, including its diet and dispersal abilities is currently underway in Sabah. The threat posed by illegal hunting of this species, and the potential trade routes remain unclear, and should be the focus of research throughout its range.|
Bernard, H. Baking, E.L., Matsubayasi, M., and Ahmad, A.H. 2012. Records of Bornean felids in and around Tabin Wildlife Reserve, Sabah, Malaysia. . Cat News 56: 4-7.
Brodie, J. and Giordano, A. J. 2012. Density of the Vulnerable Sunda clouded leopard Neofelis diardi in a protected area in Sabah, Malaysian Borneo. . Oryx 46: 427-430.
Brodie, J. F., Giordano, A. J., Zipkin, E. F., Bernard, H., Mohd-Azlan, J. and Ambu. L. 2015. Correlation and persistence of hunting and logging impacts on tropical rainforest mammals. . Conservation Biology 29: 110-121.
Buckley-Beason, V.A., Johnson, W.E., Nash, W.G., Stanyon, R., Menninger, J.C., Driscoll, C.A., Howard, J., Bush, M., Page, J. E., Roelke, M. E., Stone, G., Martelli, P. P., Wen, C., Ling, L., Duraisingam, R.K., Lam, P.V. and O'Brien, S.J. 2006. Molecular Evidence for Species-Level Distinctions in Clouded Leopards. Current Biology 16: 2371-2376.
Cheyne, S.M. and Macdonald, D.W. 2011. Wild felid diversity and activity patterns in Sabangau peat-swamp forest, Indonesian Borneo. Oryx 45: 119-124.
Cheyne S.M., Sastrimidjaja, W.J., Muhalir, Rayadin Y., Limin, S.H. and Macdonald D.W. In prep. Mammalian Communities as indicators of disturbance across Indonesian Borneo.
Cheyne, S. M., Stark, D. J., Limin, S. H., & Macdonald, D. W. 2013. First estimates of population ecology and threats to Sunda clouded leopards Neofelis diardi in a peat-swamp forest, Indonesia. Endangered Species Research 22: 1-9.
Christiansen, P. 2008. Species distinction and evolutionary differences in the clouded leopard (Neofelis nebulosa) and Diard’s clouded leopard (Neofelis diardi). . Journal of Mammology 89: 1435–1446.
Christiansen, P. 2009. Neotype of Neofelis diardi (Mammalia: Carnivora: Felidae). . Zootaxa 2110: 58-68.
Cuvier, G. 1823. Recherches sur les Ossements Fossiles oú l’on rétablit les Charactéres de plusieurs Animaux d’ont les Revolutions du Globe one détruit les Espéces. IV. . Les Ruminans et les Carnassiers Fossiles, Dufour et Dócagne, , Paris.
Gaveau, D.L., Sloan, S., Molidena, E., Yaen, H., Sheil, D., Abram, N.K., Ancrenaz, M., Nasi, R., Quinones, M., Wielaard. N., and Meijaard, E. 2014. Four decades of forest persistence, clearance and logging on Borneo. PLoS ONE 9(7).
Gordon, C. H. and Stewart, A.-M. E. 2007. The use of logging roads by clouded leopards. Cat News 47: 12-13.
Hearn, A.J., Ross, J., Macdonald, D.W., Bolongon, G., Cheyne, S.M., Azlan, M., Samejima, H., Brodie, J., Alfred, R., Boonratana, R., Meiri, S., Rustam, Bernard, H., Loken, B., Augeri, D.M., Heydon, M., Hon, J., Liat, L.B., Mathai, J., Nakabayashi, M., Pilgrim, J.D., Bezujien, M., Hall, J., Lammertink, M., Meijaard, E., Breitenmoser-Würsten, C., Kramer-Schadt, S., and Wilting, A. In prep. Predicted distribution of the Sunda clouded leopard Neofelis diardi in Borneo.
Hearn, A. J., Ross, J., Pamin, D., Bernard, H., Hunter. L., & Macdonald, D. W. 2013. Insights into the spatial and temporal ecology of the Sunda clouded leopard Neofelis diardi. Raffles Bulletin of Zoology 61(2): 871-875.
Holden, J. 2001. Small cats in Kerinci Seblat National Park, Sumatra, Indonesia. Cat News 35: 11-14.
IUCN. 2015. The IUCN Red List of Threatened Species. Version 2015-4. Available at: www.iucnredlist.org. (Accessed: 19 November 2015).
Kitchener, A.C., Beaumont, M.A. and Richardson, D. 2006. Geographical Variation in the Clouded Leopard, Neofelis nebulosa, Reveals Two Species. Current Biology 16: 2377-2383.
Kitchener, A.C., Richardson, D. and Beaumont, M.A. 2007. A new old clouded leopard. Cat News 46: 26-27.
Macdonald, E.A., Cushman, S.A., Landguth, E.L., Hearn, A.J., Malhi, Y., and Macdonald, D.W. In prep B. Clouded leopard population connectivity and gene flow in the landscape: the past, present and future.
Macdonald, E.A., Cushman, S.A., Landguth, E.L., Malhi, Y., and Macdonald, D.W. In prep A. Patterns and predictions of deforestation risk in Borneo.
Matsuda, I., Tuuga, A., & Higashi, S. 2008. Clouded leopard (Neofelis diardi) predation on proboscis monkeys (Nasalis larvatus) in Sabah, Malaysia. Primates 49: 227-231.
McCarthy, J. L., Wibisono, H. T., McCarthy, K. P., Fuller, T. K., & Andayani, N. 2015. Assessing the distribution and habitat use of four felid species in Bukit Barisan Selatan National Park, Sumatra, Indonesia. . Global Ecology and Conservation 3: 210-221.
Meijaard, E. 2004. Biogeographic history of the javan leopard Panthera pardus based on a craniometric analysis. Journal of Mammalogy 85: 302-310.
Miettinen, J., Shi, C., Tan, W. J., & Liew, S. C. 2012. 2010 land cover map of insular Southeast Asia in 250-m spatial resolution. Remote Sensing Letters 3: 11-20.
Mohamed, A., Samejima, H., and Wilting, A. 2009. Records of five Bornean cat species from Deramakot Forest Reserve in Sabah, Malaysia. . Cat News 51: 14-17.
Morino, L. 2010. Clouded leopard predation on a wild juvenile siamang. Folia Primatologica 81: 362-368.
Nájera, F., Bolongon, G., Abram, N., Goossens, B., Ambu, L., Macdonald, D.W, and Hearn, A.J. 2013. Observation of a road-killed Sunda clouded leopard in Malaysian Borneo. . Cat News 58: 42-43.
Pacifici, M., Santini, L., Di Marco, M., Baisero, D., Francucci, L., Grottolo Marasini, G., Visconti, P., Rondinini, C. 2013. Generation length for mammals. Nature Conservation 5: 87-94.
Pusparini, W., Wibisono, H.T., Reddy, G.V., Tarmizi, T. and Bharata, P. 2014. Small and medium sized cats in Gunung Leuser National Park, Sumatra, Indonesia. Cat News Special Issue(8): 4-9.
Rabinowitz, A. R., Andau, P. and Chai, P. P. K. 1987. The Clouded leopard in Malaysian Borneo. Oryx 21(2): 107.
Rautner, M. 2005. Borneo: treasure island at risk. WWF Germany, Frankfurt, Germany.
Ridout, M. S., & Linkie, M. 2009. Estimating overlap of daily activity patterns from camera trap data. Journal of Agricultural, Biological, and Environmental Statistics 14(3): 322-337.
Ross, J., Hearn, A. J., Bernard, H., Secoy, K., & Macdonald, D. W. 2010. A framework for a Wild Cat Action Plan for Sabah. Global Canopy Programme, Oxford, UK.
Ross, J., Hearn, A. J., Johnson, P. J., & Macdonald, D. W. 2013. Activity patterns and temporal avoidance by prey in response to Sunda clouded leopard predation risk. . Journal of Zoology 290(2): 96-106.
Sollmann, R., Linkie, M., Haidir, I. A., & Macdonald, D. W. 2014. Bringing clarity to the clouded leopard Neofelis diardi: first density estimates from Sumatra. Oryx 48(4): 536-539.
Wilting A., Buckley-Beason, V.A., Feldhaar, H., Gadau, J., O’Brien, S.J. and Linsenmair, S.E. 2007. Clouded leopard phylogeny revisited: support for species and subspecies recognition. Frontiers in Zoology 4: 15.
Wilting A., Buckley-Beason, V. A., Feldhaar, H., Gadau, J., O’Brien, S. J., & Linsenmair, K. E. 2007. Clouded leopard phylogeny revisited: support for species recognition and population division between Borneo and Sumatra. . Frontiers in Zoology 4: Article 15 (10 pp.).
Wilting, A., Christiansen, P., Kitchener, A. C., Kemp, Y. J. M., Ambu, L., & Fickel, J. 2011. 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. Molecular Phylogenetics and Evolution 58: 317-328.
Wilting, A., Fischer, F., Bakar, S. A. and Linsenmair, K. E. 2006. Clouded leopards, the secretive top-carnivore of South-East Asian rainforests: Their distribution, status and conservation needs in Sabah, Malaysia. BMC Ecology 6: 16.
Wilting, A., Mohamed, A., Ambu, L. N., Lagan, P., Mannan, S., Hofer, H., & Sollmann, R. 2012. Sunda Clouded Leopard Neofelis diardi density in two used forests in Sabah, Malaysian Borneo. . Oryx 46: 423-426.
Yeager, C. P. 1991. Possible antipredator behaviour associated with river crossings by proboscis monkeys (Nasalis larvatus). American Journal of Primatology 24: 61-66.
Yue, S., Brodie, J. F., Zipkin, E. F., & Bernard, H. In press. Oil palm plantations fail to support mammal diversity. . Ecological Applications..
|Citation:||Hearn, A., Ross, J., Brodie, J., Cheyne, S., Haidir, I.A., Loken, B., Mathai, J., Wilting, A. & McCarthy, J. 2015. Neofelis diardi. The IUCN Red List of Threatened Species 2015: e.T136603A50664601. . Downloaded on 27 May 2016.|
|Feedback:||If you see any errors or have any questions or suggestions on what is shown on this page, please provide us with feedback so that we can correct or extend the information provided|