Overview

Brief Summary

Biology

Like the mainland clouded leopard, Diard's clouded leopard has impressive tree climbing abilities, and is capable of running head-first down tree trunks, climbing about on the underside of branches, and hanging upside down by its hind feet with the tail providing balance. The ability to climb trees allows it to forage for food in the canopy although it mainly uses the tree branches for resting (5). Interestingly, it appears to spend less time in the trees in regions where tigers and leopards are absent (1). It was originally thought that the long canines were for preying on large ungulates (7), though recent studies show that this species feeds on a variety of terrestrial and arboreal prey including proboscis monkeys, grey leaf monkeys, young sambar deer, barking deer, mouse deer, bearded pigs, palm civets, fish and porcupines (1). Prey is either stalked on the ground or ambushed from above (6) (9). Clouded leopards are believed to be solitary, except when breeding or accompanied by cubs (10). Most information about clouded leopard reproduction comes from captive individuals (6), but as these generally comprise the mainland species, very little is known about the reproductive biology of Diard's clouded leopard (3).
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Description

Previously considered to be a subspecies of the clouded leopard (Neofelis nebulosa), Diard's clouded leopard has recently been recognised as a distinct species. Aside from genetic and anatomical differences (3) (4), Diard's clouded leopard can be recognised by its darker, grey or greyish-yellow fur and smaller cloud-like markings (3). These markings, from which the common name is derived, comprise ellipses partially edged in black, with the insides a darker colour than the background colour of the pelt (5). The limbs and underbelly are marked with large black ovals, and the back of the neck is conspicuously marked with two black bars (6). The thickly-furred tail is exceptionally long, often equivalent to the body length, and is boldly marked with black rings (5). Well adapted to forest life, Diard's clouded leopard has stout legs and broad paws, which make it excellent at climbing trees and creeping through thick forest (2). Perhaps the most remarkable feature of clouded leopards is that, in proportion to their body size, they possess the largest canines of all the cats (7) and Diard's clouded leopard has, on average, even larger and more knife-like canines than Neofelis nebulosa (4). Indeed, although they are considered to be of an unrelated evolutionary lineage, clouded leopards have independently evolved teeth and jaws that are remarkably similar to the primitive members of the extinct group of sabretoothed cats, such as the 8-10 million year-old, puma-sized Paramachairodus from Europe and Asia (8).
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Distribution

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;Christiansen2009; 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 km. Data regarding SundaClouded Leopardhabitat 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 km, and thus a range-wide total AOO of 451,900 km. 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).
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Sundaland clouded leopards occur on the islands of Borneo and Sumatra in the Malay Archipelago. It is currently unknown if Sundaland clouded leopards are on Batu, a smaller island close to Sumatra. Fossils of clouded leopards have been found on the island of Java. Sundaland clouded leopards are believed to have diverged from mainland clouded leopards approximately 1.5 million years ago due to geographic barriers. The presence of Sundaland clouded leopards on the Malay Peninsula has not been confirmed.

Biogeographic Regions: oriental (Native )

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Range

Diard's clouded leopard is believed to be restricted to the islands of Sumatra and Borneo (1).
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Physical Description

Morphology

Sundaland clouded leopards are medium-sized and have large spots along its entire body which resemble clouds. Their spots are darker and larger than the mainland clouded leopard, and the coat is darker than that of their mainland counterpart. The spots on their coat are outlined in black and the inside is darker than their primary coat color. They have two distinct black bars on the back of their necks, as well as large black ovals on the venter. The exceptionally long tail, which helps with balance while traveling in the canopy, is covered in thick fur and has a number of dark black rings along its length. They have short legs and broad paws, which make it exceptional at climbing trees, as well as moving silently through dense forests. The hind feet have very flexible joints, which allow them to descend from the canopy head first. Their flexible joints also enables them to hang from a branch using only their hind feet while using their forefeet to capture prey. Sundaland clouded leopards have exceptionally large canine teeth, which can be up to 5 cm long. In porportion to their body size, they have the largest canines of any felid. The morphology of their jaws and teeth are similar to those of extinct saber-toothed cats. Head and body length ranges from 60 to 110cm, tail length ranges from 55 to 91cm long and weight ranges from 15 to 30 kg. Males are generally larger than females.

Range mass: 15 to 30 kg.

Range length: 115 to 201 cm.

Other Physical Features: endothermic ; bilateral symmetry

Sexual Dimorphism: male larger

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Ecology

Habitat

Habitat and Ecology

Systems
  • Terrestrial
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Habitat and Ecology

Habitat and Ecology
The SundaClouded Leopardappears 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 SundaClouded Leopardthrough 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 SundaClouded 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 Leopardin 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 Leopardto 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 (Njera et al. 2013, Hearn et al. unpublished data).

On Sumatra, the SundaClouded Leopardwas 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 SundaClouded Leopard.

Little is known regarding the spatial ecology of the SundaClouded Leopard. A female SundaClouded Leopard, residing in a selectively logged forest in Sabah, occupied a home-range of 16.1 km and a core-range of 5.4 km (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 SundaClouded Leopardon 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 thatClouded Leopardsmay 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 SundaClouded 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 SundaClouded 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 SundaClouded Leopards 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 SundaClouded Leopardshunting primates during the day (Davis 1962, Matsuda et al. 2008, Morino 2010).

Systems
  • Terrestrial
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Sundaland clouded leopards are primarily forest-dwellers, however, they have been observed in other habitats as well. They are most abundant in hilly areas on the island of Sumatra, and have been observed in the lowland rain forests of Borneo as well, below 1500 m. Evidence suggests that they occupy low-elevation habitats due to the absence of large predators such as tigers. They are often sighted on the periphery of logged forests and close to human civilizations, likely due to extensive habitat loss occurring throughout its geographic range. Sundaland clouded leopards are about six times more abundant on Borneo than on Sumatra. They are highly arboreal and are particularly fond of trees overhanging ridges or cliffs. In areas containing tigers, a known predator of Sundaland clouded leopards, they rarely descend to the ground and are thought to travel through the canopy. They appear to be more arboreal on the island of Sumatra than in other areas of their geographic range, possibly due to sympatry with tigers. Despite their highly arboreal nature, they occasionally travel alongside logging roads as well.

Range elevation: < 1500 (low) m.

Habitat Regions: tropical ; terrestrial

Terrestrial Biomes: forest

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A forest-dwelling species, Diard's clouded leopard is most abundant in hilly, montane areas of rainforest on Sumatra, but may also be found in lowland rainforest on Borneo. Small numbers may occur in areas of logged forest and around the outskirts of oil-palm plantations (1).
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Trophic Strategy

Sundaland clouded leopards are carnivorous and feed on a wide variety terrestrial and arboreal prey. They regularly feed on sambar deer, barking deer, mouse deer, bearded pig, Palm civet, gray leaf monkey, fish, birds and porcupines. They have been observed preying upon proboscis monkeys as well; specifically, they target infant proboscis monkeys or juvenile females. They are ambush predators and attack prey from the canopy. They have been known to remove the limbs of their prey and bring them into trees for protection against leeches and to relax while feeding.

Animal Foods: birds; mammals; fish

Primary Diet: carnivore (Eats terrestrial vertebrates)

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Associations

There is no information available regarding the potential impact that Neofelis diardi has on its local environment.

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The Sundaland clouded leopard is a large predator and has very few predators of its own. They are illegally hunted by humans for their coats as well certain body parts that are used in traditional medicine. On Sumatra, tigers are thought to be important predators, however, this has not been confirmed. During the day, Sundaland clouded leopards remain in the canopy more than during the night, presumably to avoid tigers. They are very well camouflaged, which likely helps reduce risk of predation.

Known Predators:

  • Humans (Homo sapiens)

Anti-predator Adaptations: cryptic

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Life History and Behavior

Behavior

There is little information available regarding communication and perception in Sundaland clouded leopards. With the exception of breeding season and when females are with cubs, they are highly solitary. They are territorial and appear to use logging roads as boundaries, which are openly and frequently crossed. They are thought to demarcate territorial bounderies with urine. There is no information available regarding intraspecific communication with mates and young.

Communication Channels: chemical

Other Communication Modes: scent marks

Perception Channels: visual ; tactile ; acoustic ; chemical

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Life Cycle

There is no information available regarding the development of Neofelis diardi.

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Life Expectancy

There is no information available regarding the average lifespan of Neofelis diardi.

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Reproduction

Neofelis diardi is thought to be seasonally monogamous. There is no further information available regarding the mating system of this species.

Mating System: monogamous

There is little information available regarding breeding behavior in Sundaland clouded leopards, and all available information was gathered by observing captive individuals. Captive breeding has been mostly unsuccessful due to aggression between mates, which occasionally results in the death of the female. If introduced at a young age, aggression is not as pronounced and has allowed for more successful breeding. They are believed to exhibit similar breeding behaviors as mainland clouded leopards. Most Sundaland clouded leopards become sexually mature around 2 years of age. Mating can occur during any month of the year, but peaks between December and March. Gestation ranges from 85 to 95 days and results in 1 to 5 cubs, with an average of 2 cubs per litter. Cubs are usually independent once they reach 10 months old and become reproductively mature by 2 years of age. The females are able to produce a litter every year.

Breeding interval: Sundaland clouded leopards breed once yearly

Breeding season: Breeding activity in Sundaland clouded leopards occurs year-round but peaks from December through March.

Range number of offspring: 1 to 5.

Average number of offspring: 2.

Range gestation period: 85 to 95 days.

Average time to independence: 10 months.

Average age at sexual or reproductive maturity (female): 2 years.

Average age at sexual or reproductive maturity (male): 2 years.

Key Reproductive Features: iteroparous ; seasonal breeding ; gonochoric/gonochoristic/dioecious (sexes separate); sexual ; fertilization (Internal ); viviparous

There is little information available regarding parental care in Sundaland clouded leopards. Mothers nurse cubs until about 10 months of age, at which time they become independent.

Parental Investment: precocial ; female parental care ; pre-weaning/fledging (Provisioning: Female)

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Conservation

Conservation Status

IUCN Red List Assessment


Red List Category
EN
Endangered

Red List Criteria
C2a(i)

Version
3.1

Year Assessed
2008

Assessor/s
Sunarto, Sanderson, J. & Wilting, A.

Reviewer/s
Nowell, K., Breitenmoser-Wursten, C., Breitenmoser, U. (Cat Red List Authority) & Schipper, J. (Global Mammal Assessment Team)

Contributor/s

Justification
Santiapillai and Ashby (1988) found confirmed evidence for clouded leopards in six areas comprising just 3% of Sumatra's land area. Density in one of these areas (the Tesso Nilo - Bukit Tigapuluh Conservation Landscape) was estimated at 1.6 adults per 100 km by camera trapping (Hutujulu et al. 2007), much lower than densities found on Borneo, probably in part due to its sympatry with the larger Sumatran tiger, whereas on Borneo there are no larger felid competitors. Clouded leopards are forest-dependent, and a continuing decline is suspected due to high deforestation rates, occurring outside protected areas (3.2-5.9%/yr: Achard et al. 2002, FWI/GFW 2001, Uryu et al. 2007), but also, to a lesser extent, inside protected areas (Gaveau et al. 2007, Kinnaird et al. 2003, Linkie et al. 2004). With a fragmented, low density population, the effective population size of the Sumatran clouded leopard is probably less than 2,500 mature individuals, with no subpopulation having an effective population size greater than 250 mature individuals (IUCN Cats Red List Workshop 2007).
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IUCN Red List Assessment


Red List Category
VU
Vulnerable

Red List Criteria
A2c; C1

Version
3.1

Year Assessed
2015

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-Wrsten, C., Lanz, T. & Breitenmoser, U.

Contributor/s
Sanderson, J. & Sunarto, S.

Justification
The elusive SundaClouded Leopardis 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 SundaClouded Leopardappears to be adaptable, and is found in a range of forest types, elevations and levels of disturbance. Despite the species apparent habitat plasticity, the SundaClouded Leopardis 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 (Njera et al. 2013, Hearn et al. unpublished data). The forests of Borneo and Sumatra are undergoing some of the worlds 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 SundaClouded Leopardon 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 km (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 km. Additional camera trap surveys on Borneo, specifically undertaken to detect and estimate the density of this felid, failed to find any evidence of the SundaClouded 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 SundaClouded Leopardpresence across the forested landscape of Borneo is somewhat patchy. The population density of the SundaClouded Leopardon 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 km (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 SundaClouded Leopards current area of occupancy (AOO) on Borneo is approximately 378,900 km (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 km, providing a total AOO of 451,900 km. 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 SundaClouded Leopardacross its AOO may be as low as 1 individual / 100 km. 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 SundaClouded Leopardprobably 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, SundaClouded Leopardsare prone to untargeted snaring.

We therefore have a strong basis upon which to infer that the total SundaClouded Leopardpopulation 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 SundaClouded Leopardgenerations, Pacifici et al. 2013) indicating a categorisation of Vulnerable under both VU C1 and VU A2c.

History
  • Vulnerable (VU)
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Sundaland clouded leopards are classified as vulnerable on the IUCN's Red List of Threatened Species. Major threats include rapid and extensive deforestation for agricultural expansion (e.g., oil palm) and settlements. Rapid deforestation to establish oil-palm plantations is a major road block to the longterm persistence of this species. Deforestation laws are rarely enforced and even wildlife sanctuaries and national forests have been somewhat deforested since 1970. Deforestation not only decreases the amount of available habitat for this species, but reduces available habitat for potential prey as well. Additional threats include illegal hunting and accidental trapping. Two subspecies have been recognized, Neofelis diardi borneensis and Neofelis diardi diardi, both of which are classified as endangered on the IUCN's Red List of Threatened Species. Sundaland clouded leopards occur in a number of protected areas throughout its geographic range and is listed under Appendix 1 by CITES (the Convention on International Trade in Endangered Species of Wild Fauna and Flora).

CITES: appendix i

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Status

Classified as Vulnerable (VU) on the IUCN Red List and is listed on Appendix I of CITES (1). Subspecies: Neofelis diardi borneensis and Neofelis diardi diardi are both listed as Endangered (EN) on the IUCN Red List (1).
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Population

Population Trend
Decreasing
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Population

Population
Our understanding of the SundaClouded Leopards 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 km, with 6 of 10 surveys providing estimates of under 1.9 ind. / 100 km. 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 km: Brodie and Giordano 2012, Danum Valley Conservation Area, 1.76 ind. / 100 km: Hearn, Ross and Macdonald unpublished data), upland (Tawau Hills National Park, 1.85 ind. / 100 km: Hearn, Ross and Macdonald unpublished data) and submontane/montane (Crocker Range National Park, 1.4 ind. / 100 km: 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 km: Wilting et al. 2012, Ulu Segama Forest Reserve, 2.55 ind. / 100 km, Tabin Wildlife Reserve, 2.26 ind. / 100 km (Hearn, Ross and Macdonald unpublished data). In Kalimantan, there are only two studies that have yielded population density estimates of the SundaClouded 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 km). 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 SundaClouded Leopardsresiding within the Sabangau peat swamp, a disturbed peat swamp forest in Central Kalimantan (0.7-4.4 ind. / 100 km). Additional SundaClouded Leopardfocused surveys on Borneo failed to find any evidence of this species, including the relatively small (ca. 43 km) 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 SundaClouded Leopardpresence 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 km (Sollmann et al. 2014).

A Maximum Entropy based habitat suitability analysis for Borneo, discussed in detail above, provided a total AOO of 451,900 km (Borneo: 378,900 km, Sumatra: 73,000 km). 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 SundaClouded Leopardacross its AOO may plausibly be as low as 1 individual / 100 km. 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.

Population Trend
Decreasing
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Threats

Major Threats
Habitat loss due to commercial logging and conversion to oil palm plantations and the increasing threat from poaching pose the greatest threats to the SundaClouded 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 (Rautner2005). 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 Borneos forests (an area of 11,167 km) 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 SundaClouded Leopardbut 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 SundaClouded Leopardnumbers and also a significant reduction in the potential gene flow between populations. Increased fragmentation of Clouded Leopardhabitat 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 SundaClouded Leopards(Njera 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).
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The principal threat to Diard's clouded leopard is the catastrophic level of deforestation that is occurring on Sumatra and Borneo as a result of logging and clearance for oil palm plantations. In addition, accidental capture in snare traps used to catch other animals, as well as deliberate poaching of this species' commercially valuable pelt are having a significant impact on its population (1).
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Management

Conservation Actions

Conservation Actions
The SundaClouded Leopardis 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 SundaClouded Leopardhas been the subject of increased research in recent years, but it remains one of the least studied of the worlds 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 SundaClouded Leopards 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.
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Conservation

Diard's clouded leopard receives national protection in Sumatra, Kalimantan, Sabah, Sarawak and Brunei, as well as international protection through its listing on Appendix I of the Convention on International Trade in Endangered Species (CITES) (1). It also occurs in most protected areas along the Sumatran mountain spine, and in the majority of protected areas on Borneo. Nevertheless, given the severity of habitat loss within this species' range, as well as the illegal hunting that is occurring, increased protective measures are urgently required (1). The recognition of Diard's clouded leopard as a distinct species has important conservation implications, as unlike the mainland clouded leopard, there is not currently a captive breeding program in place (3). Hopefully, given its new status, efforts will be made to establish a healthy captive population.
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Relevance to Humans and Ecosystems

Benefits

Sundaland clouded leopards occasionally prey on livestock from villages surrounded by vast forest in Sumatra and Borneo. There are no records of Sundaland clouded leopards attacking humans.

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Sundaland clouded leopards are illegally hunted for their coats and various body parts are used in traditional medicine. Tissue samples from carcasses have been used in phylogenetic research, which has helped establish the relationship of this species to other felids.

Positive Impacts: body parts are source of valuable material; source of medicine or drug

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Wikipedia

Sunda clouded leopard

The Sunda clouded leopard (Neofelis diardi), also known as the Sundaland clouded leopard, is a medium-sized wild cat found in Borneo and Sumatra. In 2006, it was classified as a separate species, distinct from its continental relative Neofelis nebulosa.[2][3]

In 2008, the IUCN classified the species as Vulnerable, with a total effective population size suspected to be fewer than 10,000 mature individuals, and a decreasing population trend.[1]

Previously, the species was known as the Bornean clouded leopard — a name publicised by the WWF in March 2007, quoting Dr. Stephen O'Brien of the U.S. National Cancer Institute as saying, "Genetic research results clearly indicate that the clouded leopard of Borneo should be considered a separate species".[4]

Characteristics[edit]

The Sunda clouded leopard is the largest cat in Borneo, and has a stocky build, weighing around 12 to 25 kg (26 to 55 lb). The canine teeth are 2 in (5.1 cm) long, which, in proportion to the skull length, are longer than those of any other extant cat. Its tail can grow to be as long as its body, aiding balance.

Its coat is marked with irregularly-shaped, dark-edged ovals which are said to be shaped like clouds, hence its common name. Though scientists have known of its existence since the early 19th century, it was positively identified as being a distinct species in its own right in 2006, having long been believed to be a subspecies of the mainland clouded leopard (Neofelis nebulosa).[2][3]

Distribution and habitat[edit]

Sunda clouded leopards are probably restricted to the Indonesian islands of Borneo and Sumatra. In Borneo, they occur in lowland rainforest, and at lower density, in logged forest. Records in Borneo are below 1,500 m (4,900 ft). In Sumatra, they appear to be more abundant in hilly, montane areas. It is unknown if there are still Sunda clouded leopards on the small Batu Islands close to Sumatra.[1]

Between March and August 2005, tracks of clouded leopards were recorded during field research in the Tabin Wildlife Reserve in Sabah. The population size in the 56 km2 (22 sq mi) research area was estimated to be five individuals, based on a capture-recapture analysis of four confirmed animals differentiated by their tracks. The density was estimated at eight to 17 individuals per 100 km2 (39 sq mi). The population in Sabah is roughly estimated at 1,500–3,200 individuals, with only 275–585 of them living in totally protected reserves that are large enough to hold a long-term viable population of more than 50 individuals.[5]

On Sumatra, Sunda clouded leopards occur most probably in much lower densities than on Borneo. One explanation for this lower density of about 1.29 individuals per 100 km2 (39 sq mi) might be that on Sumatra clouded leopards co-occur sympatrically with the tiger, whereas on Borneo clouded leopards are the largest carnivores.[6]

The first documentary video of a Sundaland clouded leopard was taken in June 2009 by a group of wildlife biologists in Sabah.[7]

Clouded leopard fossils have been found on Java, where it perhaps became extinct in the Holocene.[8]

Ecology and behaviour[edit]

The habits of the Sunda clouded leopard are largely unknown because of the animal's secretive nature. It is assumed that it is generally solitary. It hunts mainly on the ground and uses its climbing skills to hide from dangers.[citation needed]

Evolutionary and taxonomic history[edit]

Illustration published 1834 in William Jardine's The Natural History of The Feline

The species was named Felis diardi in honor of the French naturalist and explorer Pierre-Médard Diard by Georges Cuvier in 1823, based on a drawing and skin allegedly from Java.[9] In the 19th century Felis diardii designated both clouded and Sunda clouded leopards, colloquially "Diard's Cat".[10]

The Sunda clouded leopard was long regarded as a subspecies of the clouded leopard, and named Neofelis nebulosa diardi. The genetic analysis of hair samples of Neofelis nebulosa and Neofelis diardi implies that they diverged 1.4 million years ago, after having used a now submerged land bridge to reach Borneo and Sumatra from mainland Asia.[2]

Results of a morphometric analysis of the pelages of fifty-seven clouded leopards sampled throughout the genus' wide geographical range indicated that there are two distinct morphological groups, differing primarily in the size of their cloud markings.[3] DNA samples from the Bornean and mainland Asia populations were used in molecular genetic analyses, revealing differences in mtDNA, nuclear DNA sequences, microsatellite and cytogenetic variation. Thirty-six fixed mitochondrial and nuclear nucleotide differences, and 20 microsatellite loci with nonoverlapping allele-size ranges distinguished the populations — a degree of differentiation equivalent to, or greater than, comparable measures among the panthera species — and strongly support a species-level distinction between Neofelis nebulosa and Neofelis diardi.[2]

In December 2006, the genus Neofelis was reclassified into two distinct species:[2][3]

Molecular, craniomandibular and dental analysis indicates subspecifical distinction of Bornean and Sumatran clouded leopards into two populations with separate evolutionary histories — a Bornean subspecies Neofelis diardi borneensis and a Sumatran subspecies Neofelis diardi diardi. Both populations are estimated to have diverged from each other during the Middle to Late Pleistocene. The split of Neofelis diardi subspecies corresponds roughly with the catastrophic super-eruption of the Toba Volcano in Sumatra 69,000–77,000 years ago. A probable scenario is that Sunda clouded leopards from Borneo recolonized Sumatra during periods of low sea levels in the Pleistocene, and were later separated from their source population by rising sea levels.[11]

Threats[edit]

Deforestation in Sumatra

Sunda clouded leopards being strongly arboreal are forest-dependent, and are increasingly threatened by habitat destruction following deforestation in Indonesia as well as in Malaysia.[1] Since the early 1970s, much of the forest cover has been cleared in southern Sumatra, in particular lowland tropical evergreen forest. Fragmentation of forest stands and agricultural encroachments have rendered wildlife particularly vulnerable to human pressure.[12] Borneo has one of the world's highest deforestation rates. While in the mid-1980s forests still covered nearly three quarters of the island, by 2005 only 52% of Borneo was still forested. Both forests and land make way for human settlement. Illegal trade in wildlife is a widely spread practice.[13]

The population status of Sunda clouded leopards in Sumatra and Borneo has been estimated to decrease due to forest loss, forest conversion, illegal logging, encroachment, and possibly hunting. In Borneo, forest fires pose an additional threat, particularly in Kaltim and in the Sebangau National Park.[14]

There have been reports of poaching of Sunda clouded leopards in Brunei's Belait District where locals are selling their pelts at a lucrative price.[15]

Conservation[edit]

Neofelis diardi is listed on CITES Appendix I, and is fully protected in Sumatra, Kalimantan, Sabah, Sarawak and Brunei. Sunda clouded leopards occur in most protected areas along the Sumatran mountain spine, and in most protected areas on Borneo.[1]

Since November 2006, the Bornean Wild Cat and Clouded Leopard Project based in the Danum Valley Conservation Area and the Tabin Wildlife Reserve aims to study the behaviour and ecology of the five species of Bornean wild cat — bay cat, flat-headed cat, marbled cat, leopard cat, and Sunda clouded leopard — and their prey, with a focus on the clouded leopard; investigate the effects of habitat alteration; increase awareness of the Bornean wild cats and their conservation needs, using the clouded leopard as a flagship species; and investigate threats to the Bornean wild cats from hunting and trade in Sabah.[16]

The Sundaland clouded leopard is one of the focal cats of the project Conservation of Carnivores in Sabah based in northeastern Borneo since July 2008. The project team evaluates the consequences of different forms of forest exploitation for the abundance and density of felids in three commercially used forest reserves. They intend to assess the conservation needs of these felids and develop species specific conservation action plans together with other researchers and all local stakeholders.[17]

Etymology[edit]

The scientific name of the genus Neofelis is a composite of the Greek word νεο- meaning "new", and the Latin word feles meaning "cat", so it literally means "new cat."[18][19]

Local names[edit]

The Indonesian common name for the clouded leopard rimau-dahan means "tree tiger" or "branch tiger".[20]

See also[edit]

References[edit]

  1. ^ a b c d e Hearn, A., Sanderson, J., Ross, J., Wilting, A., Sunarto, S. (2008). "Neofelis diardi". IUCN Red List of Threatened Species. Version 2014.3. International Union for Conservation of Nature. 
  2. ^ a b c d e 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., Wen, C., Ling, L.; Duraisingam, R.K., Lam, V.P., O'Brien, S.J. (2006). "Molecular Evidence for Species-Level Distinctions in Clouded Leopards". Current Biology 16 (23): 2371–2376. doi:10.1016/j.cub.2006.08.066. PMID 17141620. 
  3. ^ a b c d Kitchener, A. C., Beaumont, M. A., Richardson, D. (2006). "Geographical Variation in the Clouded Leopard, Neofelis nebulosa, Reveals Two Species". Current Biology 16 (23): 2377–2383. doi:10.1016/j.cub.2006.10.066. PMID 17141621. 
  4. ^ "New Species Declared: Clouded Leopard found on Borneo and Sumatra". ScienceDaily. 2007. Retrieved 26 November 2010. 
  5. ^ Wilting, A., Fischer, F., Abu Bakar, S. 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. doi:10.1186/1472-6785-6-16. PMC 1654139. PMID 17092347. 
  6. ^ Hutujulu, B., Sunarto, Klenzendorf, S., Supriatna, J., Budiman, A. and Yahya, A. (2007). Study on the ecological characteristics of clouded leopard in Riau, Sumatra. In: J. Hughes and M. Mercer (eds.) Felid Biology and Conservation: Programme and Abstracts : An International Conference, 17–20 September 2007, Oxford. Oxford University, Wildlife Conservation Research Unit
  7. ^ Mohamed, A. and Wilting, A. (2009). Sundaland clouded leopard Neofelis diardi. Filmed at Deramakot Forest Reserve, Sabah, Malaysia. Conservation of Carnivores in Sabah
  8. ^ Meijaard, E. (2004). Biogeographic history of the Javan leopard Panthera pardus based on a craniometric analysis. Journal of Mammalogy 85: 302–310.
  9. ^ Cuvier, G. (1823). Recherches sur les ossemens fossiles; ou, l'on retablit les caracteres de plusiers animaux dont les revolutions du globe ont detruit les especes. Les Ruminans et les Carnassiers Fossiles, Volume IV. Paris: G. Dufour & E. d'Ocagne
  10. ^ Ripley, G. (1858). The New American Cyclopedia. D. Appleton and Company. p. 543. 
  11. ^ Wilting A., Christiansen P., Kitchener A. C., Kemp Y. J. M., Ambu L., Fickel, J. (2010). "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 (2): 317–28. doi:10.1016/j.ympev.2010.11.007. PMID 21074625. 
  12. ^ Gaveaua, D. L. A., Wandonoc, H., Setiabudid, F. (2007). "Three decades of deforestation in southwest Sumatra: Have protected areas halted forest loss and logging, and promoted re-growth?". Biological Conservation 134 (4): 495–504. doi:10.1016/j.biocon.2006.08.035. 
  13. ^ Rautner, M., Hardiono, M., Alfred, R. J. (2005), Borneo: treasure island at risk. Status of Forest, Wildlife, and related Threats on the Island of Borneo, WWF Germany 
  14. ^ Povey, K., Sunarto, H. J.G., Priatna, D., Ngoprasert, D., Reed, D., Wilting, A., Lynam, A., Haidai, I., Long, B., Johnson, A., Cheyne, S., Breitenmoser, C., Holzer, K., Byers, O. (eds.) CBSG. (2009), Clouded Leopard and Small Felid Conservation Summit Final Report, IUCN/SSC Conservation Breeding Specialist Group: Apple Valley, MN. 
  15. ^ Shahminan, F., Begawan, B. S. (2010). Poaching threatens clouded leopards The Brunei Times, 19 December 2010.
  16. ^ Hearn, A., Ross, J. (2006). "Bornean Wild Cat and Clouded Leopard Project". Cat Project of the Month – November 2006. IUCN/SSC Cat Specialist Group. 
  17. ^ Wilting, A., Mohamed, A. (2009). "Consequences of different forest management strategies for felids in Sabah, Malaysia". Cat Project of the Month – May 2009. IUCN/SSC Cat Specialist Group. 
  18. ^ Perseus Digital Library. Greek Dictionary νεο Headword Search Result
  19. ^ Perseus Digital Library. Latin Dictionary feles Headword Search Result
  20. ^ Horsfield, T. (1825). Description of the Rimau-Dahan of the inhabitants of Sumatra, a new species of Felis, discovered in the forests of Bencoolen, by Sir Thomas Stamford Raffles, late Lieutenant-Governor of Fort Marlborough. Zoological Journal of London 1: 542−554.
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