Carnivora (Wilson & Reeder, 2005)

Corlett, Richard T., 2011, Vertebrate Carnivores And Predation In The Oriental (Indomalayan) Region, Raffles Bulletin of Zoology 59 (2), pp. 325-360 : 338-345

Carnivora

Felidae . – The cats are hypercarnivores, requiring a higher proportion of protein in their diets than most other mammals ( Sunquist & Sunquist, 2009) and apparently unable to detect the sweetness of carbohydrates ( Li et al., 2005). They are highly specialised for killing live prey, with none of the morphological compromises required by the more omnivorous diet of many other Carnivora . Hypercarnivory is associated with specialisation of the teeth for slicing, rather than crushing, along with other shared features of the skull and dentition that we recognise as “cat-like”, although other groups of carnivores have occupied this niche in the past ( Van Valkenburgh, 1988). Oriental felids range in size from the 1-kg rusty-spotted cat ( Felis rubiginosus ) to the 75–260-kg tiger ( Panthera tigris ) and the similar sized, but much more restricted, lion ( Panthera leo ). Note that generic assignments in much of the regional literature do not reflect the current understanding of their relationships (Agnarsson et al., 2010).

The five largest extant Oriental felids, the tiger ( Panthera tigris ), lion ( P. leo ), leopard ( P. pardus ), snow leopard ( P. uncia ) and clouded leopards ( Neofelis spp. ), together with the New World jaguar ( P. onca ), form a well-supported monophyletic group ( Johnson et al., 2006). The tiger and leopard were historically almost ubiquitous, while the lion was confined to northwest India, the clouded leopard to the eastern half of the Region, and the snow leopard to the northern margins of the Region, mostly above the treeline ( Sunquist & Sunquist, 2009).

Siberian tigers are the largest living felids, but tropical tigers are similar in size to African lions, with adult males 100–260 kg and adult females 75–160 kg in the Region ( Sunquist & Sunquist, 2009). Tigers from the Sunda shelf islands (Sumatra and, until recently, Java and Bali) are considerably smaller than those from northern India and southern China. Tigers occupied a variety of more or less wooded habitats throughout the Region, except on the islands of Sulawesi and the Philippines, and Borneo, where they may have survived into the last millennium (Cranbrook, 2010). Primary lowland rainforests appear to be a relatively unfavourable habitat for tigers, with very low population densities compared with habitats where a higher primary productivity at ground level supports a higher density of potential prey (Kawanishi & Sunquist, 2004). The tiger is the most specialised felid and densities at diverse sites are predicted fairly accurately by the densities of the ungulate prey on which they depend ( Karanth et al., 2004; Kawanishi & Sunquist, 2004). Tigers select large prey, with mean prey weights above 50 kg at most sites and prey> 100 kg forming an important component of the diet wherever available (Biswas & Sankar, 2002; Bagchi et al., 2003; Joseph & Thomas, 2006; Andheria et al., 2007; Avinandan et al., 2008; Sunquist & Sunquist, 2009). Deer—particularly Axis axis (70 kg) and Rusa (Cervus) unicolor (180 kg)—and wild pigs (<120 kg) were the major prey at most study sites, but they also prey regularly on gaur Bos frontalis (450–900 kg), where it is available, as well as a huge range of other species, from frogs, birds and fish to sun bears and the calves of elephants and rhinos. Adult elephants and rhinoceroses escape predation by tigers, and tapirs (395 kg) seem to be avoided for unknown reasons (Kawanishi & Sunquist, 2004). Tigers kill more people than any other carnivore in the Region. The predictive model of Karanth et al. (2004) assumes that each tiger consumes around 50 ungulates a year, representing 10% of all available prey. In general, tiger populations do not appear to survive for long in the absence of large prey, but the major prey species were <50 kg in Srisailam Tiger Reserve in southern India ( Reddy et al., 2004) and were <20 kg Muntiacus muntjak in the Huai Kha Khaeng Wildlife Sanctuary in Thailand ( Rabinowitz, 1989).

Through most of their range, tigers coexist with leopards and the dhole ( Cuon alpinus ). Tigers are socially dominant and sometimes kill leopards, but large dhole packs have been known to chase and, occasionally, kill tigers. There is considerable overlap in the types of prey taken by the three species, but dholes are strictly diurnal, while tigers are mostly crepuscular or nocturnal, and leopards take a wider range of generally smaller prey (Joseph & Thomas, 2006; Andheria et al., 2007; Ahmed & Khan, 2008; Wang & Macdonald, 2009; Sunquist & Sunquist, 2009). The historical range of tigers also overlapped in the west of the Region with that of the Asian subspecies of lion, Panthera leo persica . Lions prefer more open habitats than tigers, but both can utilise the mosaic of grassland and open forest which supports the highest ungulate biomass in Asia. Individual male lions have apparently killed tigers in captivity (according to a range of uncheckable Internet sources), and male lions in South Africa regularly take prey more than three times their own weight (Radloff & Du Toit, 2004), which tigers do not. Moreover, unlike all other wild felids, lions normally hunt in groups (1–15, with a mean of three in Asian lions ( Singh 1995)), which must make them the most formidable predators in the Region. African lions weigh 85–225 kg but there appear to be no reliable weights for the Asian subspecies. They are now confined to dry deciduous forest in and around the Gir National Park in northwest India, where the last 400 or so individuals prey largely on adults of Axis axis , Rusa unicolor , Boselaphus tragocamelus (120–240 kg), wild pigs, and domestic cattle and buffalo ( Meena et al., 2011).

The most widespread felid, the leopard (29–70 kg in Asia), was absent only from Sumatra, Borneo, Sulawesi and the Philippines, as well as true deserts and alpine areas on the mainland. Not surprisingly for a species with such a wide geographic and environmental range, and which must compete almost everywhere with one larger felid and several smaller ones, adaptability is its most striking characteristic. Oriental leopards hunt mostly on the ground, eating whatever they can catch, but they climb well and monkeys are important prey, particularly where ungulates are rare (Joseph & Thomas, 2006; Sunquist & Sunquist, 2009; Wang & Macdonald, 2009). Most prey weigh 5–45 kg, including juveniles of larger species, but they can apparently kill much larger animals (c. <200 kg) and can survive on smaller prey (Arivazhagan et al., 2005; Odden & Wegge, 2009). In Africa, the preferred prey mass is c. 25 kg ( Hayward et al., 2006). Leopards readily eat carrion and it is not always clear whether large species recorded in scats were killed or not. In Sanjay Gandhi National Park, on the edge of Mumbai, dogs and rodents were the commonest items in scats (Edgaonkar & Chellam, 2002).

The snow leopard is confined to high mountains from Pakistan to China. Slightly smaller than the leopard (22–52 kg), it is a rock specialist, living mostly above the treeline, but it moves lower in winter and in some regions year round ( Sunquist & Sunquist, 2009). Over much of its range, the blue sheep or bharal, Pseudois nayaur (45 kg), is the most important prey, along with the ibex, Capra siberica (130 kg), Himalayan tahr, Hemitragus jemlahicus (130–180 kg), other ungulates, domestic stock (including 250-kg horses, yaks and cattle), and smaller animals, such as marmots, hares and pheasants (Gao, 1987; Sunquist & Sunquist, 2002; Bagchi & Mishra, 2006; Lovari et al., 2009). They reportedly chase prey further than other big cats.

The clouded leopards are considerably smaller (10-25 kg) than the other members of this group and are also the species about which least is known. Two species are currently recognised: N. nebulosa in mainland Southeast Asia and South China and N. diardi on the islands of Borneo and Sumatra ( Sunquist & Sunquist, 2009). Both are largely confined to dense evergreen forests. The relatively small size, long tail, short legs and broad paws have been considered adaptations to an arboreal life. Most sightings have been on the ground, but recorded prey includes arboreal squirrels, slow lorises, macaques, leaf monkeys, proboscis monkeys and orangutans, as well as a range of medium-sized terrestrial species, such as hog deer (35 kg), muntjac (<25 kg), mouse deer, pigs, pangolins and civets ( Grassman et al., 2005a; Matsuda et al., 2008; Sunquist & Sunquist, 2009). Clouded leopards have the largest canine teeth in relation to body size of any living felid and the widest maximum gape angle (c. 85 o), but the significance of this is unknown.

The Asiatic golden cat, Catopuma temminckii (8–16 kg), occurs through much of the eastern Oriental Region, from northeast India to southern China and south to Sumatra, but not Borneo ( Sunquist & Sunquist, 2002). The smaller but otherwise similar bay cat, C. badia (2–4 kg), occurs only on Borneo, where it appears to be widespread but rare ( Kitchener et al., 2004). Very little is known about the ecology of the golden cat, despite its wide range, and nothing at all about the bay cat. Golden cats appear to be mostly terrestrial, although they climb well. Two adults radio-tracked in Thailand ranged over 33–48 km 2 ( Grassman et al., 2005a) and both this study and a camera-trapping study in Malaysia (Mohd. Azlan & Sharma, 2006) showed no clear diurnal pattern of activity. The diet includes a variety of small vertebrates up to the size of mouse deer and the dusky leaf monkey ( Trachypithecus obscurus ), although there are reports of larger prey, including domestic sheep, goats and water buffalo calves (Gao, 1987; Lim, 2002; Sunquist & Sunquist, 2009). The marbled cat ( Pardofelis marmorata ) (2–5 kg) has a wide distribution in closed forests from northeast India to southwest China, and south to Borneo and Sumatra, but very little is known about its ecology. The big feet and very long tail, and its behaviour in captivity, are consistent with the marbled cat being more arboreal than most other Oriental felids. Activity seems to be largely nocturnal and crepuscular ( Grassman et al., 2005a). Scattered reports on its diet mention rats, birds and squirrels ( Sunquist & Sunquist, 2009).

The caracal ( Caracal caracal ) had a similar historical range to the cheetah, but is now rare in Pakistan and on the verge of extinction in India ( Sunquist & Sunquist, 2002). It is a slender, long-legged cat that can sprint short distances at high speed and jump 2 m into the air from a standing start to catch a bird. Over the whole geographical range, body weights are 6–20 kg ( Sunquist & Sunquist, 2009), but Indian caracals are apparently much smaller than those from Africa and the Middle East, at around 6 kg ( Mukherjee et al., 2004). In the Region it inhabited a range of habitats from semi-desert to open dry forest. Caracals hunt mainly at night on the ground, but also climb well. Sunquist & Sunquist (2002) suggest they are “gazelle cats”, specialised to hunt the 15–30 kg gazelles found throughout their historical range, but smaller prey dominates recorded diets. In Rajasthan, west India, 84% of scats (n = 25) had rodent remains, 36% birds, 20% reptiles and 40% invertebrates ( Mukherjee et al., 2004). Each caracal was estimated to eat 8–9 rodents (each weighing 16–77 g) per day. Caracals, like cheetahs, were kept by Indian princes for hunting, and were apparently trained to hunt terrestrial prey up to the size of the barasingha ( Cervus duvauceli , 145 kg) and nilgai ( Boselaphus tragocamelus , 170 kg), as well as arboreal squirrels (Divyabhanusinh, 2002).

As well as being the only carnivore species lost from the Region in historical times, the cheetah, Acinonyx jubatus , a member of the otherwise American puma lineage, is isolated phylogenetically, morphologically and ecologically from other Oriental felids. It is a largely diurnal predator that catches most of its prey in short, very high-speed chases (<102 km hr- 1 in Africa) ( Sunquist & Sunquist, 2009). Approximately the same weight as a leopard (22–65 kg in Africa), it is taller, slimmer and morphologically specialised for running, with an elbow joint more similar to a canid or hyena than other felids (Andersson, 2004). Cheetahs use the claw of the first digit of the front paw to hook fleeing prey off balance, in contrast to other large felids that knock the prey off balance ( Londei, 2000). Four hundred years ago, cheetahs were common in open and semi-open habitats in central and western India, but the last known Indian cheetahs were shot in 1947 ( Sunquist & Sunquist, 2002). The subspecies still survives in Iran. Their major prey seems to have been the blackbuck, Antilope cervicapra (38 kg), and most other recorded prey are in a similar size range (e.g., Jebeer gazelle ( Gazella bennettii ), wild sheep, Ovis orientalis , and Persian ibex, Capra aegagrus , in central Iran; Farhadinia & Hemami, 2010). Indian princes used trained cheetahs to hunt and even imported African ones when they became scarce in India (Divyabhanusinh, 2002).

The jungle cat, Felis chaus , is the largest (3–12 kg) and most widespread of this genus in the Region, ranging from Pakistan to Sri Lanka and southwest China, but not Malaysia or Indonesia ( Sunquist & Sunquist, 2009). Despite its name, this species is not usually found in dense forest, preferring grasslands, open forests and cultivated areas, often, but not always, near water. It hunts mostly on the ground and is often seen during the day. One was clocked at 32 km hr 1 from a car ( Roberts, 2005). Most reported prey is <1 kg, although they occasionally kill larger prey such as juvenile ungulates. A study in Rajasthan, western India, found that 73% of 69 scats contained the remains of rodents, 42% birds, 26% reptiles and 23% invertebrates ( Mukherjee et al., 2004). Each cat was estimated to eat 3–5 rodents (weighing 16–77 g) a day. Most scats in cultivated areas in Pakistan were also dominated by rodents (Khan & Beg, 1986). Fish and frogs are eaten elsewhere.

The Asian form of the wild cat, Felis silvestris (3–4 kg), enters the Region only in Pakistan and northwest India, where it is associated mostly with semi-desert habitats ( Sunquist & Sunquist, 2009). It is largely terrestrial and nocturnal, feeding mostly on rodents, but also taking occasional larger prey (<3–4 kg) such as young ungulates, as well as birds, reptiles and invertebrates. It hybridises freely with the closely related domestic cat. The sand cat, F. margarita (2–3 kg), has an even more restricted range within the Region, occurring only in sandy deserts in western Pakistan ( Roberts, 2005). It is a desert specialist, in terms of its morphology and physiology, its highly developed hearing, and its opportunistic diet, including small mammals, birds, reptiles and invertebrates.

Free-ranging domestic cats, Felis catus , occur in and around most human settlements in the Region, but the distribution and abundance of genuinely feral populations have not been investigated. Cats were domesticated later than dogs and most still supplement their diet by hunting, so one would expect the domestic-feral transition to be relatively easier. Feral cats on Iriomote Island (24 oN) were associated with garbage dumps, but also preyed on rats, birds, frogs and invertebrates ( Watanabe et al., 2003). On nearby Okinawa (27 oN), 11 feral cat scats contained the remains of birds, shrews, snakes, frogs and invertebrates ( Kawauchi & Sasaki, 2002). On Hahajima (26 oN), in the Bonin Islands, feral cats apparently prey on seabirds, particularly the wedge-tailed shearwater ( Puffinus pacificus ), as well as rats (Kawakami & Fujita, 2004), while a free-ranging domestic cat concentrated on small birds, particularly the introduced Zosterops japonicus (Kawakami & Higuchi, 2002) . Elsewhere in the Region, feral cats prey on both egg-laying megapodes and newly emerged chicks on the Moluccan island of Haruka ( Heij, 2001). Feral cats have been responsible for a large percentage of global vertebrate extinctions, particularly on islands ( Nogales et al., 2004), so more work is clearly needed on the potential threat posed by this species in the Region. Domestic cats are assumed to have less impact per cat (Kays & DeWan, 2004), but can live at unnaturally high densities and in areas that cannot support feral cat populations.

The leopard cat, Prionailurus bengalensis (1–4 kg), has the widest Oriental distribution of any small felid and is absent only from the extreme west and from the Philippines and Sulawesi ( Sunquist & Sunquist, 2002). Like all members of this genus, leopard cats are excellent swimmers and the species occurs on many offshore islands ( Watanabe, 2009). A distinctive subspecies, P. bengalensis iriomotensis , occurs on the southern Ryukyu island of Iriomote (24 oN). Leopard cats occupy a wide range of mostly forested habitats, but also seem to survive in deforested, human-dominated landscapes better than most other Asian felids. Although usually considered to be nocturnal, radio-telemetry and camera-trapping studies have found them to also be active during the day ( Rabinowitz, 1990; Grassman et al., 2005b; Mohd. Azlan & Sharma, 2006). Rats and mice dominate the diet in most studies, but they have also been reported to take a wide variety of other small vertebrates, including squirrels, tree shrews, hares, mouse deer ( Tragulus javanicus , 3 kg), birds, lizards, snakes, frogs and fish (Gao, 1987; Rabinowitz, 1990; Khan, 2004; Grassman et al., 2005b; Austin et al,. 2007; Rajaratnam et al., 2007; Sunquist & Sunquist, 2009). Gao (1987) says they will attack Muntiacus . The well-studied Iriomote subspecies (3–5 kg) has an extremely broad diet, including rats, the Ryukyu flying fox, birds, skinks, snakes, frogs and invertebrates (Nowell & Jackson, 1996; Izawa et al., 2000).

The fishing cat ( Prionailurus viverrinus ) and the flat-headed cat ( P. planiceps ) have a lot in common (Nowell & Jackson, 1996; Sunquist & Sunquist, 2009). Both have an elongated head, short legs and a very short tail, contributing to their rather un-catlike appearances, with the fishing cat most often compared to a civet (hence “ viverrinus ”) and the flat-headed cat to a mustelid. Both are associated largely with wetlands, both take to water readily, and both prey heavily on fish. Both species use their forepaws to search for prey underwater ( Iwaniuk et al., 2001). The flat-headed cat, however, is much more specialised as a fish-eater, with its more completely webbed toes and a long and pointed second upper pre-molar, which enables it to grasp slippery prey. Other peculiar features of this small (1.5–2.5 kg) cat, including its flattened head, large close-set eyes, and small ears, may also be adaptations to its semi-aquatic existence. The little information on its natural diet shows it to be dominated by fish, plus other aquatic animals, including frogs and crustacea, although flat-headed cats have been known to prey on poultry. It is confined to peninsular Thailand, the Malay Peninsula, Sumatra and Borneo. The fishing cat, in contrast, is a larger (5–16 kg), less specialised cat, with a wider but discontinuous distribution that includes parts of Pakistan, Sri Lanka, southwest and northeast India, Nepal, Bangladesh, and Southeast Asia to Sumatra and Java, but not Borneo or the Malay Peninsula. Fish appear to be its most frequent prey at most sites, but it is also known to take birds, rodents, a small Indian civet ( Viverricula indica ), snakes, other reptiles, frogs and a variety of domestic animals, including dogs ( Sunquist & Sunquist, 2009).

The rusty spotted cat, Prionailurus rubiginosus , may be the smallest cat species, at 1.0– 1.6 kg ( Sunquist & Sunquist, 2009). It is found in Sri Lanka and much of peninsular India, but little is known about its ecology. It occurs in a range of habitats from grassland to dense forest, is mostly nocturnal, and seems to hunt mostly on the ground, consuming a variety of small vertebrates.

Prionodontidae . – The Asian linsangs (two species of Prionodon ) have traditionally been included among the civets, but molecular data strongly support family status (Agnarsson et al., 2010). This is consistent with their cat-like appearance and behaviour, retractile claws, and hypercarnivorous dentition, as well as their largely carnivorous diet. Both species are smaller than any cat or Oriental civet, with the banded linsang ( P. linsang ) weighing 600–800 g and the spotted linsang ( P. pardicolor ) around 600–1200 g ( Gaubert, 2009). The spotted linsang occurs in hill forests from Nepal and northeast India, through northern Myanmar and Thailand, to Laos, Vietnam and southwest China, while the banded linsang occurs in forests from southern Myanmar and Thailand to Sumatra, Borneo and Java. Both species appear to be equally at home in trees and on the ground and are reported to feed on rats, squirrels, birds, lizards, snakes, frogs and insects (Gao, 1987; Van Rompaey, 1993; Shrestha, 1997; Nowak, 1999; Gaubert, 2009). Like cats but unlike most civets, they do not seem to eat fruit.

Hyaenidae . – Hyenas are characterised by a bone-cracking dentition, with relatively wide premolars ( Van Valkenburgh, 1988). The striped hyena ( Hyaena hyaena ), was widespread in suitable habitats from Pakistan to Nepal and Bangladesh, and south through India to the Nilgiri Hills (Gurung & Singh, 1996). The bone-cracking niche is currently unoccupied in the east of the Region and in densely forested areas of the west, presumably because large carcasses are too rare or too difficult to locate in closed forest. However, the spotted hyena ( Crocuta crocuta ; 45–80 kg) survived in southern China until at least the Late Pleistocene and a giant hyena ( Pachycrocuta brevirostris ;> 100 kg) was present in nonforest habitats in the Region earlier in the Pleistocene (Corlett, 2010). Oriental hyenas weigh 30–50 kg and occur in a wide range of more or less open habitats, where they are usually solitary foragers ( Prater, 1980; Shreshta, 1997; Roberts, 1997; Nowak, 1999). They seem to live largely as scavengers and their ability to crush the largest bones enables them to exploit carcasses already picked clean by vultures ( Roberts, 1997). There are also reports of striped hyenas appropriating kills from leopards ( Prater, 1980; Roberts, 1997). However, striped hyenas also kill a variety of small and medium-sized mammals, including domestic sheep, goats and dogs, and the occasional human child. They are also known to eat tortoises and considerable quantities of fruit. Twenty-six hyena scats collected in Rajasthan, northwest India, contained the remains of Axis axis (35% of scats), Boselaphus tragocamelus (14%), domestic cattle (17%) and goats (14%), and hares, Lepus nigricollis (7%), as well as fruit, a bird and a rodent, but it was not possible to distinguish predation from scavenging (Sankar & Jethwa, 2002).

Herpestidae . – Almost every habitat in the Region supports at least one (<3) species of mongoose ( Herpestes ). All eight or so Asian mongoose species have long, slender bodies, short legs with non-retractile claws, a tapering snout, small rounded ears and a long tail. The smallest species, such as H. javanicus , weigh <1.2 kg while the largest ( H. urva and H. vitticollis ) can weigh> 3 kg ( Prater, 1980; Lekagul & McNeely, 1988; Roberts, 1997). All are predominantly terrestrial, but some species climb trees and some are strongly associated with water. Most species appear to be largely diurnal, although literature reports are not consistent on this. All species are usually solitary foragers and, although pairs are sometimes reported, there is no evidence of cooperative hunting.

Asian mongooses are opportunistic predators of large invertebrates and small vertebrates, but there have been no detailed studies of the diet of wild populations within their native range. Invertebrates are the most frequent items in scats, but small vertebrates are probably more important for most species in terms of biomass consumed (e.g. Gorman, 1975; Abe et al., 1999). Herpestes javanicus (often called H. auropunctatus ) is by far the best-studied species, but almost entirely in areas where it has been introduced to control rats or snakes outside its native range, often with disastrous consequences for endemic vertebrates ( Roy, 2002; Quinn & Whisson, 2005). One such well-studied introduction was to Amami Island (28 oN), on the northeast fringes of the Region, where 30 individuals were released in 1979 (Abe et al., 1999; Yamada, 2002; Watari et al., 2008). On this island, the vertebrate component of the diet includes mammals (particularly Rattus tanezumi ), birds, frogs and reptiles, but very rarely the snake ( Protobothrops flavoviridis ) it was intended to control. Eight percent of scats contained remains of the endangered endemic rabbit, Pentalagus furnessi , which has an adult weight of 2–3 kg. Generally similar diets have been reported for other introduced and native populations of this species ( Gorman, 1975; Prater, 1980; Gao, 1987; Lekagul & McNeely, 1988; Roberts, 1997; Ogura et al., 2002).

Scattered information for the other species suggests that they share broadly similar diets as well as an ability to kill prey larger than themselves ( Phillips, 1980; Prater, 1980; Lekagul & McNeely, 1988; Roberts, 1997). An ability to kill highly venomous snakes has also been widely reported, particularly for H. edwardsi , and seems to reflect their speed and agility together with a relative resistance to both hemorrhagic and neurotoxic venoms (Dunham, 2003). One of the two largest species, H. vitticollis , has been reported to prey on hares ( Lepus nigricollis , 2–6 kg) and mouse deer ( Tragulus meminna , 3–4 kg) ( Phillips, 1980; Van Rompaey & Jayakumar, 2003). The diet of the other large species, H. urva , is dominated by aquatic vertebrates and invertebrates, according to most accounts (Lekagul & McNeely, 1988; Chuang & Lee, 1997; Van Rompaey, 2001), but Wang (1999) found that the remains of rodents and snakes were common in scats from southern China.

Viverridae . – After the exclusion of the linsangs, most Asian civets fall into three groups—true civets (Viverrinae), palm civets (Paradoxurinae) and the rest (Hemigalinae). Most Asian civets are omnivores, consuming vertebrates, invertebrates and plant foods, particularly fruits (Corlett, 1998; Jennings & Veron, 2009). This is reflected in their dentition, which is much less specialised than that of the felids. The vertebrate prey of civets is generally small in comparison with felids of similar body mass. As many as six civet species can coexist in the same area (e.g. Mohd. Azlan, 2006).

The four Oriental species of Viverrinae are nocturnal, terrestrial (at least when foraging), solitary omnivores. The smallest and most widespread species, Viverricula indica (2–4 kg), occupies almost all habitat types, from rainforest to semidesert, but appears to be most common in non-forest or partly forested habitats, often near human habitations. Reported diets include varying proportions of small vertebrates (rodents, shrews, birds and reptiles), invertebrates and fruit ( Prater, 1980; Gao, 1987; Lekagul & McNeely, 1988; Roberts, 1997; Chuang & Lee, 1997; Wang & Fuller, 2003a; Jennings & Veron, 2009). The three Viverra species are larger (3–11 kg), but have essentially similar diets, with rodents the most consistent component (Wemmer & Watling, 1986; Gao, 1987; Lekagul & McNeely, 1988; Auffenberg, 1988; Lim, 1991; Colón, 1999; Jennings & Veron, 2009). Jha (1999) says that V. zibetha in India hunts prey larger than itself, including dogs, goats, deer and even buffalo calves, but I can find no other support for this statement.

The Paradoxurinae are largely nocturnal and generally more arboreal and more frugivorous than the Viverrinae. The most widespread species, Paradoxurus hermaphroditus (2–5 kg), is largely arboreal in forest, but has adapted to life in humanmade structures in and around human settlements (Lekagul & McNeely, 1988; Krishnakumar & Balakrishnan, 2003). It appears to prefer fruit when this is available, but can live on invertebrates or small vertebrates (mammals, birds and reptiles) when fruit is rare (Gao, 1987; Auffenberg, 1988; Joshi et al., 1995). The other three species of Paradoxurus are largely arboreal, forest frugivores, but P. lignicolor , endemic to the Mentawai Islands, is reported to prey on village chickens (Abegg, 2003). Paguma larvata (3–5 kg) is less arboreal and less exclusively nocturnal than Paradoxurus , but has a similar diet, at least seasonally dominated by fruit, and also thrives near human settlements in some areas (Gao, 1987; Lekagul & McNeely, 1998; Wang & Fuller, 2001, 2003a; Dudgeon & Corlett, 2004).

Macrogalidia musschenbroeki is the only native species in the order Carnivora on Sulawesi, although two other viverrids have been introduced. It inhabits forests from the lowlands to the montane zone, but also takes poultry from villages (Wemmer & Watling, 1986). Although a skilful climber, it probably feeds mostly on the ground. Its diet is apparently dominated by fruits, especially palm fruits, but it also eats rodents, the nocturnal dwarf cuscus ( Strigocuscus celebensis ), and birds, and local people say it takes piglets of Sus celebensis and, occasionally, domestic pigs. The binturong, Arctictis binturong , is a large (9–20 kg) arboreal species of dense Southeast Asian forests, with slow and deliberate movements. It and the Neotropical kinkajou ( Potos ) are the only Carnivora with truly prehensile tails. Very little is known of its diet but it appears to be largely frugivorous. Arctogalidia trivirgata (2–2.5 kg) is morphologically distinct and its relationships require further study (Agnarrson et al., 2010). It is a nocturnal, arboreal species of forest canopies in Southeast Asia, which is reported to feed on squirrels, birds, frogs, insects and fruits, although no quantitative dietary information is available (Lekagul & McNeely, 1988; Jennings & Veron, 2009).

The Hemigalinae includes four distinctive-looking civets of Southeast Asian forests, all of which seem to feed largely on invertebrates, although vertebrate prey may be taken occasionally ( Nowak, 1999): Hemigalus derbyanus (1–3 kg), Diplogale hosei (1.4 kg), Chrotogale owstoni (2–3.5 kg) and the otter civets, Cynogale bennettii (4 kg).

Canidae . – Most canids are adapted to relatively open habitats, so most species are confined to the west and north of the Region. The Oriental dogs span the full range of variation found in the family, from solitary omnivores to pack-hunting predators of animals considerably larger than themselves.

Two similar-sized pack-hunting taxa, the dhole ( Cuon alpinus ) and the wolf ( Canis lupus , in the broad sense) effectively partitioned the Region in the past, although they coexist in habitat mosaics today. Oriental dholes occupy most of the forested parts of India and southern China, south through Southeast Asia to Sumatra and Java. Males are 15–20 kg, females 10–12 kg and reported pack size averages around 3– 12 adults, with an extreme of 40 (Cohen, 1978; Gao, 1987; Indrawan et al., 1996; Karanth & Sunquist, 2000; Silbero- Zubiri, 2009). Dholes are diurnal hunters and are most active in the early morning and evening. They specialise on hunting whatever ungulates are locally available, but are also reported to take hares, rodents and other small vertebrates at some sites (Cohen et al., 1978; Prater, 1980; Barnett et al., 1981; Fox, 1984; Gao, 1987; Venkataraman et al., 1995; Indrawan et al., 1996; Wang, 1999; Karanth & Sunquist, 2000; Austin, 2002; Grassman et al., 2005c; Joseph & Thomas, 2006; Silbero-Zubiri, 2009). Unlike wolves, dholes rarely attack domestic stock, although they are reported to take goats in Kerala (Veermani et al., 1996) and sheep in Bhutan (Wang & Macdonald, 2009). Most chases do not last long (<500 m). Most prey in India is <50 kg, but dholes regularly kill adult male Axis axis (75 kg) and Rusa unicolor , including adult males (> 200 kg), is the major prey species at some sites (Austin, 2002; Kumara et al., 2004; Grassman et al., 2005c; Joseph & Thomas, 2006; Borah et al., 2009). Dholes in Java kill adult female Rusa timorensis (95 kg) and sub-adult or female Bos javanicus (<500 kg), the latter in packs of 9-23 individuals ( Indrawan et al., 1996). Conversely, the small (<4) packs of dhole at a rainforest site in Peninsula Malaysia preyed mostly on mouse deer (2–4.5 kg) (Kawanishi & Sunquist, 2008). Large prey usually dies from shock or loss of blood as a result of multiple injuries. Jog et al. (2005) found that the density of sarcocysts of the protozoan parasite Sarcocystis in the heart muscles of dhole-killed Axis axis was significantly higher than in animals that died of other causes, suggesting that dholes select infected prey and, since the dhole is an obligatory host for the sexual cycle of the parasite, that the relationship between predator and parasite could be mutualistic. Dholes appear to have substantially larger area requirements for persistence in protected areas than do tigers, perhaps because their larger home ranges (70 km 2 for dholes vs. 17 km 2 for tigers in India) bring them more often into conflicts with people (Woodroffe & Ginsberg, 1998). Meat-stealing from dhole kills by local people is a problem in some areas ( Kumara et al., 2004).

Wolves once occupied most habitats in the Region other than tropical forests and deserts. Recent mtDNA studies identified three distinct lineages of wolves: one confined to lowland India and Pakistan, one found at high elevations from eastern Nepal and Tibet to eastern Kashmir, and the “wolf-dog” clade that includes all other wolves in the Region, as well as those in the Palaearctic and Nearctic, and all domestic and feral dogs ( Sharma et al., 2004; Aggarwal et al., 2007). Further studies are needed, but it is possible that these three lineages deserve recognition as separate species. Little is known about the ecology of the Himalayan wolf lineage, but there have been a number of studies of the lowland wolves ( Jhala, 2003; Kumar & Rahmanai, 2008). Although male wolves from India and Pakistan can attain 24– 27 kg ( Roberts, 1997; Prater 1980), most seem to be <20 kg, which is more similar to the dhole than to the wolves of the Palaearctic and Nearctic ( Nowak, 1999). Centuries of persecution because of attacks on livestock and, less commonly, children ( Shahi, 1982; Rajpurohit, 1999) have made it very difficult to reconstruct the natural predatory behaviour of lowland Indian wolves. Most wolves in human-dominated landscapes combine scavenging and predation on livestock ( Shahi, 1982; Jhala, 2003; Singh & Kumara, 2006). Away from human settlements, Oriental wolves take a variety of wild ungulates, but also hares, rodents and other small vertebrates (Kumar & Rahmani, 2000, 2008; Jethva & Jhala, 2003; Jhala, 2003). Blackbucks ( Antelope cervicapra ) are the major prey in western India, with the wolves preferring old or injured adult males (c. 40 kg) (Kumar & Rahmani, 2008). Reported pack sizes are mostly 1–7, with a maximum of 12, and most kills are made at night, although this may be partly to avoid persecution. Most chases last less than 0.5 km ( Jhala, 2003), but they can continue considereably longer (Kumar & Rahmani, 2008). Untill recently, lowland wolves would have competed with cheetahs in much of their range.

Domesticated dogs ( C. lupus familiaris ) appear to have originated from wolves by multiple independent domestication events and have probably been genetically separate from them for around 8000 years ( Verginelli et al. 2005). Today, every possible intermediate from fully domesticated to fully feral dogs can be found in the Region, mostly near human settlements (Corbet & Hill, 1992; Irion et al., 2005; Vanak & Gompper, 2009, 2010). The diversity of size, shape and other characteristics declines along this gradient and most feral dogs in the Region have converged on (or retained?) a single, dingo-like form. Little is known about the ecology of feral and semi-feral dogs in the Region. Hunting behaviour was presumably selected against during domestication, because of the threat to people and other domesticates, but feral dogs sometimes hunt wild prey larger than themselves in small packs, as well as taking small vertebrates, carrion and garbage (Auffenberg, 1981; Jhala, 1993; Kawauchi & Sasaki, 2002; Manor & Saltz, 2004; Dudgeon & Corlett, 2004; Das et al., 2006; Vanak & Gompper, 2009).

The golden jackal ( Canis aureus ) looks like a small (7–12 kg) wolf. It occurs from Thailand and Myanmar westwards, mostly in relatively open habitats and often near human settlements. Rodents, carrion and a wide range of plant material are the most frequently recorded dietary items, along with other small mammals, ground birds and their eggs, reptiles, frogs, fish and invertebrates ( Schaller, 1967; Prater, 1980; Khan & Beg, 1986; Sarker & Ameen, 1990; Roberts, 1997; Nowak, 1999; Jaeger et al., 2007; Silbero- Zubiri, 2009). They are mostly nocturnal and usually seen alone, although packs of 3–5 have also been reported. Pairs or larger groups may attack domestic goats, langurs ( Presbytis spp. ), small deer and the young of larger species ( Stanford, 1989; D’Cunha, 1996; Silbero-Zubiri, 2009).

Foxes ( Vulpes spp. ) are small (1–10 kg), relatively shortlegged canids that occupy most of the western part of the Region, as well as southern China and northern Vietnam, but are absent from the tropical forests of Southeast Asia. V. bengalensis (1.8–3.2 kg) occupies most of India, V. vulpes enters the northern part of the Region from Pakistan to South China, and the ranges of V. cana (c. 1 kg) and a desert specialist, V. rueppelli (1.5–3 kg), extend into the northwest of the Region. A highland species, V. ferrilata (3–5 kg), enters the northern margins of the Region in Nepal. Foxes live in pairs but are usually solitary hunters. Diets are usually dominated by rodents and other small mammals, but also include ground birds, reptiles, frogs, invertebrates and fruits ( Prater, 1980; Roberts, 1997; Sharma, 1997; Nowak, 1999; Home & Jhala, 2009; Vanak & Gompper, 2009; Silbero- Zubiri, 2009).

The raccoon dog ( Nyctereutes procyonoides ) (4–8 kg) enters the northeast of the Region in southern China and northern Vietnam, where it inhabits a variety of forest types. It is a nocturnal omnivore, feeding on small mammals, birds ,

reptiles, frogs, invertebrates and large amounts of fruit (Gao, 1987; Silbero-Zubiri, 2009).

Ursidae . – Four bear species more or less partitioned the Region in historical times, although their distributions have now been greatly reduced. The sloth bear ( Melursus ursinus ) occupied Sri Lanka and most of the Indian subcontinent, except deserts and high mountains. The sun bear ( Helarctos malayanus ) occupied most of Southeast Asia, from southwest China south to Sumatra and Borneo. The Asiatic black bear ( U. thibetanus ) occupied forests in southern China and a belt across the Himalayan ranges west to Pakistan. The range of this species had a broad overlap with the sun bear in continental Southeast Asia and a narrower one with the sloth bear in northern India, with all three species apparently coexisting in parts of eastern India. Finally, the brown bear ( U. arctos ) occurs largely in alpine habitats on the margins of the Region in India and Pakistan.

The Oriental bears appear to feed largely on fruit and plant material, and on invertebrates, and most evidence for carnivory is anecdotal. However, all species are capable of killing an adult human. The Asiatic black bear (males 110– 180 kg, females 59–90 kg) is largely vegetarian ( Schaller, 1969; Gao, 1987; Reid et al., 1991; Roberts, 1997; Hwang, 2003; Sathyakumar & Viswanath, 2003), but they have been reported to dig Alticola voles out of their burrows ( Johnsingh, 2003). They sometimes kill sheep and goats ( Roberts, 1997) and, occasionally, larger domestic animals up to the size of an adult buffalo ( Prater, 1980; Nowak, 1999). Presumably they also take wild vertebrates opportunistically ( Hwang, 2003), like the related American black bears. Although brown bears (<400 kg) are relatively carnivorous in some parts of their huge natural range (e.g., Xu et al., 2006), the few Oriental reports suggest that they are largely herbivorous here ( Prater, 1980; Roberts, 1997). Like black bears, they are reported to dig rodents out their burrows and some individual bears kill sheep, goats and other livestock.

The sun bear (27–65 kg) is the smallest and most arboreal bear. The long tongue and protrusible lips are presumably adaptations to insectivory, but they do not seem to maintain body condition on invertebrates alone and become almost totally frugivorous during mast fruiting periods ( Wong et al, 2004; Fredriksson et al., 2006). They are also reported to kill small vertebrates, including tortoises, and, occasionally livestock, as well as scavenging on tiger kills ( Wong et al., 2002; Ward & Kynaston, 1995; Nowak, 1999; Fitzgerald & Krausman, 2002; Wong et al., 2004). The sloth bear (males 80–150 kg, females 55–95 kg), is even more specialised as an insectivore, feeding largely on ants, termites and fruits in a wide range of habitat types (Garshelis et al., 1999; Sacco & Van Valkenburgh, 2004). Most studies have found no direct evidence of vertebrate consumption (e.g. Schaller, 1967; Laurie & Seidensticker, 1977; Gokula et al., 1995).

Mustelidae . – There are around 23 species of mustelids in the Region, although several of these barely penetrate the northern margins. The family occurs throughout, except in the Philippines (excluding Palawan) and Sulawesi, but is represented only by otters on Sri Lanka. The two species of stink badger ( Mydaus ) are now placed in the skunk family, Mephitidae (Flynn et al., 2005) . The diet of stink badgers is dominated by invertebrates, with some plant material, and there are no records of them taking vertebrates.

The weasels ( Mustela spp. ) are represented by nine species (three only marginally) that range from the Himalayan region to southern China and south to Sumatra, Borneo and Java, but the genus is absent from most of India (Corbet & Hill, 1992). The Oriental species are small (50–800 g), longbodied, short-legged carnivores that are mostly terrestrial, diurnal or nocturnal. The little dietary information available from the Region suggests they feed mostly on rodents and other small vertebrates ( Prater, 1980; Gao, 1987; Roberts, 1997, 2005; Jha, 1999; Nowak, 1999), but more detailed studies of M. sibirica in mainland China (Gao, 1987), on Taiwan ( Wu, 1999), and on the southern Palaearctic island of Tsushima (Tatara & Doi, 1994), suggest that this species, at least, is an extreme opportunist, with the diet dominated by rodents, shrews, birds, frogs, invertebrates or fruits at different sites or seasons.

The marbled polecat ( Vormela peregusna ) (600 g) inhabits open, semi-arid areas in Pakistan, where it feeds on a variety of rodents and other small vertebrates ( Roberts, 1997). It is mostly nocturnal and, although it can climb, it captures most of its prey on or under the ground. There are three species of martens ( Martes spp. ) in the Region, the widespread M. flavigula , M. gwatkinsii in southern India and M. foina on the northern fringes of the Region. M. flavigula and M. gwatkinsii are bigger than weasels (2–3.5 kg), largely diurnal or crepuscular, and have omnivorous diets. They hunt in trees and on the ground, sometimes in pairs or in larger, presumably family, groups ( Prater, 1980; Gao, 1987; Ramakantha, 1994; Roberts, 1997, 2005; Nowak, 1999; Austin & Tewes, 1999; Madhukumar, 2002; Mudappa, 2002; Grassman et al., 2006; Larivière & Jennings, 2009). Their reported prey includes a wide variety of small vertebrates, up to the size of Tragulus meminna (2.5 kg) and juveniles of larger ungulates, plus some fruit and insects.

The ferret badgers ( Melogale spp. ), which are not closely related to the true badgers, are small (1–3 kg) nocturnal animals that feed largely on invertebrates, although small mammals, birds, reptiles and frogs are also occasionally eaten ( Prater, 1980; Gao, 1987; Chuang & Lee, 1997; Nowak, 1999; Wu, 1999). The Asian badger ( Meles leucurus ) (3.5–9 kg) enters the northeast of the Region and presumably feeds on small vertebrates, invertebrates and fruit like its European relative ( M. meles ) ( Larivière & Jennings, 2009). The hog badgers ( Arctonyx spp. ) (7–14 kg) occur from the eastern Himalayas to China and south to Thailand and Sumatra. Although usually reported to feed largely on invertebrates, 43 of 45 scats attributed to hog badgers in Jiangxi, China, contained the remains of rodents, and some contained hares, hedgehogs, birds or snakes ( Wang, 1999; Wang & Fuller, 2003b). The ratel or honey badger ( Mellivora capensis ) (c. 9 kg) ranges from southern Africa to the drier parts of peninsular India. It is reported to feed on a variety of small vertebrates, invertebrates and plant material in the Region ( Prater, 1980; Roberts, 1997; Pati et al., 2001), but is known to kill larger prey in Africa ( Larivière & Jennings, 2009).

The Region’s otter species ( Aonyx cinerea , Lutra lutra , L. sumatrana , Lutrogale perspicillata ) feed primarily on aquatic prey, mostly at night, but all four species take terrestrial rodents and birds when the opportunity arises ( Prater, 1980; Gao, 1987; Kruuk et al., 1994; Kanchanasaka, 1997; Roberts, 1997; Nowak, 1999; Lariviere, 2003; Anoop & Hussain, 2005). Prater (1980) says that Lutrogale perspicillata “takes to jungle hunting like other land carnivores” when pools and streams dry out.

Ailuridae . – The red panda ( Ailurus fulgens ) feeds mostly on bamboo leaves, but it is also said to occasionally consume small rodents, young birds and bird eggs (Gao, 1987; Nowak, 1999).